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ISSN 0968-0470 



Bulletin of 

The Natural History 

Museum 



Zoology Series 



TH£ NATURAL 
HISTORY &40S6UM 



P??E$EWTTD 
Of NB*AL UBRAPti 




VOLUME 67 NUMBER 2 29 NOVEMBER 2001 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum 
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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) 



© The Natural History Museum, 2001 



ISSN 0968-0470 



Zoology Series 

Vol. 67, No. 2, pp. 109-207 



The Natural History Museum 
Cromwell Road 
London SW7 5BD 



Issued 29 November 2001 



typeset by Ann Buchan (Typesetters), Middlesex 

Printed in Great Britain by Henry Ling Ltd.. at the Dorset Press, Dorchester, Dorset 



/jxCSfoo^'l ) 



Bull. not. Hist. Mus. bond. (Zool.) 67(2): 109-136 



Issued 29 November 2001 



Type material of Stegocephalidae Dana, 1855 
(Crustacea, Amphipoda) in the collections of 
The Natural History Museum, London, 
including the description of seven new species 



T l XL NATURA L 
HISTORY MUSEUM 

tnv.uit.^ QCT 20Q1 

FHh RENTED 
GENERAL USR ARY 



J0RGEN BERGE AND WIM VADER 

Troms0 Museum, Dept. of Zoology, University of Troms0, 9037 Tromsfi, Norway, e-mail: joergenb@ 

ANTON GALAN 

Marine Research Institute, Skiilagata 4, Reykjavik, Iceland, e-mail: anton@hafro.is 



CONTENTS 



Synopsis 109 

Introduction 109 

Material and methods 1 10 

Systematics 1 10 

Andaniexis australis K.H. Barnard. 1932 110 

Andaniexis americana sp.nov 1 1 1 

Andaniexis gloriosa sp.nov 113 

Andaniexis pelagica sp.nov 1 13 

Andaniopsis africana sp.nov 117 

Bathystegocephalus globosus (Walker, 1909) 119 

Phippsia gibbosa (Sars, 1883) 120 

Phippsiella rostrata K.H. Barnard, 1932 121 

Stegocephaloides attingens K.H. Barnard. 1932 124 

Stegocephaloides australis K.H. Barnard, 1916 127 

Stegocephaloides calypsonis sp.nov 129 

Stegocephaloides boxshalli sp.nov 129 

Stegocephaloides ledoyeri sp.nov 131 

Discussion 135 

References 135 



SYNOPSIS. Six species belonging to the amphipod family Stegocephalidae Dana, 1855 (Crustacea) are redescribed and 
figured. Seven new species are also described: Andaniexis americana sp.nov., A. gloriosa sp.nov. and A. pelagica sp.nov., 
Andaniopsis africana sp.nov., Stegocephaloides boxshalli sp.nov., 5. calypsonis sp.nov. and S. ledoyeri sp.nov. The type 
material of Stegocephaloides attingens K.H. Barnard 1916 comprises two species, 5. attingens and 5. boxshalli sp.nov. 
Stegocephaloides calypsonis is formally described for the first time, but was originally described in the PhD thesis of Anton 
Galan(1984). 



INTRODUCTION 



Many species in the amphipod family Stegocephalidae Dana, 1855 
were described at the end of the 19 th or the beginning of the 20 ,h 
century. These descriptions usually took the form of a limited 
descriptive text accompanied by very few figures, typically showing 
only one or two diagnostic characters. Very few of these species 
have been later redescribed in any detail, and when they have, the 
description was usually based on new material without any refer- 
ence to the type material (e.g. Barnard, 1962, 1964; Ledoyer, 1986; 
all referring to Andaniexis australis K.H.Barnard 1932). Con- 



sequently, for a number of stegocephalid species, there is consider- 
able confusion in terms of morphology, generic and specific status. 
The vast collections of The Natural History Museum in London 
(UK) include type material of nine stegocephalid species, none of 
which has ever been redescribed from the type material. Three of 
these species belong to the genus Andaniotes Stebbing, 1897; which 
have recently been dealt with elsewhere (Berge, 2001a.). In the 
present paper, the type material of six species is redescribed and 
figured: Andaniexis australis K.H.Barnard, 1932, Bathystego- 
cephalus globosus (Walker, 1909), Phippsia gibbosa (Sars, 1883), 
Phippsiella rostrata K.H.Barnard, 1932, Stegocephaloides attingens 
K.H.Barnard, 1916 and S. australis K.H.Barnard, 1916. In addition 



© The Natural History Museum, 2001 



no 



J. BERGE, W. VADER AND A. GALAN 



to these six species, one. Stegocephaloides calypsonis sp.nov., is 
Formally described for the first time, although it was first recognized 

bj \nton Galan in his unpublished Ph.D. thesis (1984). 

\s a 'spin-off 1 effect of describing some of the above mentioned 
species, six Further new species are described: Andaniexisamericana, 
A.gloriosa and A. pelagica,Andaniopsis africana, Stegocephaloides 
boxshalli and S. ledoyeri. All these species have previously been 
assigned to one of the six redescribed species. 

The phylogenetic relationships and generic status of the species 
described herein are not discussed in the present paper, which is one 
of a series (together with Berge & Vader 1997 a-d) that will 
collectively revise all the genera and species in the family, and 
eventually lead to a phylogenetic analysis of the Stegocephalidae 
Berge & Vader. in press). 



MATERIAL AND METHODS 

The present study is based primarily on material from the collections 
of the Natural History Museum in London, UK (BMNH). Addi- 
tional material has been borrowed from the American Museum of 
Natural History, New York (AMNH) and the South African Mu- 
seum, Cape Town (SAM). 

All dissected appendages were mounted in poly vinyl-lactophenol 
and stained with rose-bengal. Figures of these appendages were 
made using a Leica compound microscope, while the habitus- 
drawings were made using a Leica dissecting microscope. Mature 
and immature females were distinguished from males by the pres- 
ence of oostegites. 

Scales on figures are all 0. 1 mm. Classification of setae and setae- 
groups follow the scheme of Berge (2001b). 

Symbols: A 1-2: Antenna 1-2; EP3: Epimeral plate 3; L: Labium; 
L*BR: Labrum; LMND: Left mandible; MX1: Maxilla 1; MX2: 
Maxilla 2; MXP: Maxilliped; Pl-7: Pereopods 1-7; PLP: palp; 
RMND: Right mandible; T: Telson; Ul-3: Uropod 1-3. 



SYSTEMATICS 



Family STEGOCEPHALIDAE Dana, 1855 
Type genus: STEGOCEPHALUS Kr0yer, 1842 

Andaniexis Stebbing, 1906 

Andania Boeck, 1871: 128. Homonym, Lepidoptera 
Andaniexis Stebbing, 1906: 94 

Andaniexis australis K.H.Barnard, 1932 

(Figs 1-3) 

Andaniexis australis K.H.Barnard, 1932: 76 
Non A. australis Barnard 1964 (= A. pelagica sp.nov.) 
Non A. australis Barnard 1962 (= A. americana sp.nov.) 
Non A. australis Ledoyer 1986 (= A. gloriosa sp.nov.) 

MATERIAL i.xamined. Syntypes 4 females, BMNH 
1936.1 1.2.588-591, 33-34°S 9-16°E, 1000m. 

Dm Rim i ion. Only the type material is known. 

Di sr KIITION 

Based on type material. Females 4— 5mm. 
Rostrum \er\ small. 




Fig. 1 Andaniexis australis, female cotype. 

Antennae short. Antenna 1 longer than antenna 2; flagellum 6- 
articulate; accessory flagellum article 2 absent. Antenna 2 peduncle 
(articles 3-5) shorter than flagellum; article 3 short, about as long as 
broad; article 4 shorter than article 5. 

Epistome produced laterally; rectangular with a long ridge on 
each side. 

Epistomal plate (medial keel) produced; small elongate medial 
ridge exceeding along the entire epistome. 

Mandible incisor transverse; incisor smooth; left lacinia mobilis 
present, reduced, distally straight, not conical. 

Maxilla 1 palp 2-articulate, oval, apex reaching beyond outer 
plate; outer plate distally rounded, ST in two parallel rows, first 
marginal and second submarginal, ST first row with 6 setae (ST 1-5, 
ST7), ST 1 ordinary (similar to ST 2—4), gap between ST5 and ST7 
present, ST A-C present and part of second row; inner plate with a 
weakly developed shoulder; setae pappose. 

Maxilla 2 ordinary; outer plate setae without distal hooks or cleft; 
inner plate setae row A covering the entire margin, clearly separated 
from row B, row A setae pappose; 3-4 first setae pectinate distally; 
row B setae proximally pappose, distally with cusps present; row C 
present; row D present, 1-3 long setae distally, setae slender. 

Maxilliped palp 4-articulate, article 2 distally unproduced, dacty- 
lus distally simple (pointed); inner plate with 1 nodular seta; medial 
setae-row present, not reduced, transverse, setae pectinate; distal 
setae-row present, setae simple; inner setae-row present, but re- 
duced to one or two setae, conspicuously large and strong; outer 
plate outer setae-row present, setae submarginal, attached normally, 
long robust, straight; inner setae-row present, well developed, setae 
long robust, slender, appressed to outer setae-row; distal setae-group 
absent. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



111 




Fig. 2 Andaniexis australis, female cotype. 

Labrum very short; lobes symmetrical and reduced. 

Labium distally broad, oval. 

Coxal plates and basis of pereopods smooth; coxae 1-3 con- 
tiguous. 

Pereopod 1 coxa not as deep as basis; propodus subovate. 

Pereopod 2 longer and thinner than pereopod 1 ; ischium elongate, 
ratio length:breadth exceeding 1.5, posterior margin with long 
plumose setae distally; propodus subovate, palm absent. 

Pereopod 4 coxa posteroventral lobe small, reaching about the 
base of the 6th pereon segment; basis anterior and posterior margins 
without long setae, with plumose setae on distal anterior margin, no 
plumose setae on distal posterior margin; ischium with plumose 
setae on posterior distal margin. 

Pereopod 6 basis posteriorly expanded, expansion conspicuous, 
rounded posteriorly, without a row of long plumose setae. 

Pereopod 7 basis anterior margin concave, distally rounded, no 
medial row of setae. 

Oostegites on pereopods 2-5. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus longer than inner. 
Uropod 2 peduncle longer than rami, outer ramus slightly longer 
than inner. Uropod 3 peduncle longer than rami, outer ramus 2- 
articulate, outer ramus equal to inner. 

Telson as long as broad, shorter than peduncle uropod 3, entire, 
apically pointed. 

Male. Unknown. 

REMARKS. This species was originally described by K.H. Barnard 



Fig. 3 Andaniexis australis, female cotype. 

in 1932, and has later been redescribed by Barnard 1962, 1964) and 
Ledoyer (1986). However, neither of these redescriptions were 
made based upon the type material. Consequently, both Andaniexis 
americana sp.nov. (see below) and Andaniexis gloriosa sp. nov. (see 
below) were wrongly identified as A. australis s.s. 

The combination of a rectangular coxa 4, a concave anterior 
margin of pereopod 7 and the presence of only one nodular seta on 
the inner plate of the maxilliped is unique for this species within the 
genus (note: the last character is unknown for A. oculatus.). From 
the description of A. oculatus, it is not possible to separated A. 
australis from A. oculatus. However, until material of the latter has 
been made available for examination, the two species are not put into 
synonomy. 

Andaniexis americana sp.nov. 

(Figs 4-5) 

Andaniexis australis Barnard 1964: 13 fig. 9 (non A. australis K.H. 
Barnard, 1932) 

Material examined. Holotype, Station V- 15-69 (see Barnard 
1964), females 3mm, 10°13'S 80°05'W, Dec. 9, 1958, 6324-6328m. 
Paratype, female 2.6mm (see Barnard 1964). 

Distribution. Only type material is known. 

Description 

Rostrum very small. 

Antennae short. Antenna 1 longer than antenna 2; flagellum 5- 
articulate; accessory flagellum article 2 present. Antenna 2 peduncle 



11. 



J. BERGE, W. VADER AND A. GALAN 




Fig. 4 Andaniexis americana sp.nov.. from: Barnard 1964: 14, fig. 9 'Andaniexis australis' 



(articles 3-5) longer than flagellum; article 3 short, about as long as 
broad; article 4 longer than article 5. 

Epistome produced laterally, rectangular with a long ridge on 
each side; epistomal plate (medial keel) produced into a small 
elongate medial ridge exceeding along the entire epistome. 

Mouthparts not elongate or pointed. 

Mandible incisor transverse; smooth; left lacinia mobilis present, 
reduced, laterally straight, not conical. 

Maxilla 1 palp 2-articulate, oval, apex reaching beyond outer 
plate; outer plate distally rounded; ST in two parallel rows, first 
marginal and second submarginal, ST first row with 6 setae (ST1-5, 
ST7), ST 1 ordinary (similar to ST 2-4), gap between ST 5 and ST 
7 present, ST A-C present and part of second row; inner plate with 
a weakly developed shoulder, setae pappose. 

Maxilla 2 ordinary; outer plate setae without distal hooks or cleft; 
inner plate setae row A covering the entire margin, clearly separated 
from row B. row A setae pappose, 3-4 first setae pectinate distally; 
row B setae proximally pappose, distally with cusps absent; row C 
present: row D present, 1-3 long setae distally, slender. 

Maxillipcd palp4-articulate. article 2 distally unproduced, dacty- 
I us distally simple (pointed): inner plate with 1 nodular setae; medial 
setae-row present, not reduced, transverse, setae pectinate; distal 
setae-row present, setae simple: inner setae-row absent; outer plate 
outer setae-row present, submarginal. setae attached normally, setae 
long robust; inner setae-row present, well developed, setae long 
robust, slender, appressed to outer setae-row; distal setae-group 
absent. 

Labium very short: lobes symmetrical and reduced. 

Labium unknown. 

il plates and basis of pcreopods smooth: coxae I -3 contiguous. 



Pereopod 1 coxa not as deep as basis; propodus subovate. 

Pereopod 2 longer and thinner than pereopod 1 ; ischium elongate, 
ratio length: breadth 

exceeding 1.5; ischium distal posterior margin with plumose 
setae; propodus subrectangular, palm absent. 

Pereopod 4 coxa posteroventral lobe small, reaching about the 
base of the 6th pereon segment; basis anterior and posterior margins 
without long setae, with plumose setae on distal anterior margin, no 
plumose setae on distal posterior margin; ischium with plumose 
setae on posterior distal margin. 

Pereopod 6 basis posteriorly expanded, expansion conspicuous, 
rounded posteriorly, without a row of long plumose setae. 

Pereopod 7 basis anterior margin concave; distally rounded; no 
medial row of setae. 

Oostegites on pereopods 2-5. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus longer than inner. 
Uropod 2 peduncle longer than rami, outer ramus as long as inner. 
Uropod 3 peduncle longer than rami, outer ramus 2-articulate. 

Telson as long as broad, shorter than peduncle uropod 3, entire; 
apically rounded, submarginal setae apically. 

Male. Unknown. 

REMARKS. This is the only recorded species of Andaniexis from 
the west coast of the American continent. The present species is a 
typical member of this genus, but is separated from all other known 
species of this genus by its very broad and powerful lacinia mobilis 
(left mandible only). As in A. australis, A. gracilis Berge & Vader, 
1997a, A. oculatus Birstein & Vinogradov, 1970 and A. stylifer 



TYPE MATERIAL OF STEGOCEPHALIDAE 



113 




Fig. 5 Andaniexis amehcana sp.nov., holotype. 

Birstein & Vinogradov, 1960, the anterior margin of pereopod 7 is 
slightly concave, while only two other known species, A. australis and 
A. gloriosa (see below), share its state of having just one nodular seta 
on inner plate of the maxilliped. Furthermore, the present species has 
coxa 4 'heart'-shaped, a character otherwise only found in A. gracilis, 
A. mimonectes Ruffo, 1975 and A. tridentata Ledoyer, 1986. 

Andaniexis gloriosa sp. nov. 

(Figs 6-7) 

Andaniexis australis Ledoyer 1986: 953-954, fig. 375. (non A. 
australis K.H. Barnard, 1932) 

Material examined. Holotype, female 7mm, st. CH90 (see 
Ledoyer 1986). Paratypes 4 females, 7-10 mm, st. CH 87 (see 
Ledoyer 1986), 3716m. Additional material, immature 2,5 mm, st. 
DE 12 (see Ledoyer 1986), 2500m. 

Distribution. Only known from the type locality (Madagascar). 

Description 

Rostrum very small. 

Antennae short. Antenna 1 shorter than antenna 2; flagellum 5- 
articulate; accessory flagellum article 2 present. Antenna 2 peduncle 
(articles 3-5) as long as flagellum; article 3 short, about as long as 
broad; article 4 shorter than article 5. 

Epistome produced laterally, rectangular with a long ridge on 
each side; epistomal plate (medial keel) produced into a small 
elongate medial ridge exceeding along the entire epistome. 



Mouthparts not elongate or pointed. 

Mandible incisor transverse; incisor smooth; left lacinia mobilis 
present, reduced, laterally straight, not conical. 

Maxilla 1 palp 2-articulate, rectangular, apex reaching beyond 
outer plate; outer plate distally rounded; ST in two parallel rows, 
first marginal and second submarginal, ST first row with 6 setae 
(ST 1-5, ST7), ST 1 ordinary (similar to ST 2-4), gap between ST 5 
and ST 7 present, ST A-C present, all part of second row; inner plate 
with a weakly developed shoulder, setae pappose. 

Maxilla 2 ordinary, outer plate setae without distal hooks or cleft; 
inner plate setae row A covering the entire margin, clearly separated 
from row B, row A setae pappose, 3-4 first setae pectinate distally; 
row B setae proximally pappose, distally without cusps; row C 
present; row D present, 1-3 long setae distally, slender. 

Maxilliped palp 4-articulate, article 2 distally unproduced, dacty- 
lus distally simple (pointed); inner plate with 1 nodular setae; medial 
setae-row absent; distal setae-row present, setae simple; inner setae- 
row present, row reduced to one or two setae, conspicuously large 
and strong; outer plate outer setae-row present, submarginal, setae 
attached normally, setae long robust, straight; inner setae-row present, 
well developed, setae long robust, slender, appressed to outer setae- 
row; distal setae-group absent. 

Labrum very short; lobes symmetrical and reduced. 

Labium distally broad, oval. 

Coxal plates and basis of pereopods smooth. Coxae 1-3 contigu- 
ous. 

Pereopod 1 coxa not as deep as basis; propodus subovate. 

Pereopod 2 longer and thinner than pereopod 1 ; ischium elongate, 
ratio length:breadth exceeding 1.5; ischium distal posterior with 
margin plumose setae; propodus subrectangular, palm absent. 

Pereopod 4 coxa posteroventral lobe small, reaching about the 
base of the 6th pereon segment; basis anterior and posterior margins 
without long setae, without plumose setae on distal anterior and 
posterior margins; ischium without plumose setae on posterior distal 
margin. 

Pereopod 6 basis posteriorly expanded, expansion conspicuous, 
rounded posteriorly; without a row of long plumose setae. 

Pereopod 7 basis anterior margin straight, distally rounded. 

Oostegites on pereopods 2-5. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus longer than inner. 
Uropod 2 peduncle longer than rami, outer ramus longer than inner. 
Uropod 3 peduncle longer than rami, outer ramus 2-articulate, outer 
ramus longer than inner. 

Telson longer than broad, shorter than peduncle uropod 3, entire, 
apically pointed, no submarginal setae on apically. 

Male. Unknown. 

Remarks. The present species is separated from all other mem- 
bers of Andaniexis by its relatively long and pointed telson. 
Furthermore, coxa 1 is relatively long and pointed (triangular), and 
is thus more similar to that of Andaniotes Stebbing, 1897. 

Unfortunately, the type material, borrowed from Museo Civico de 
Storia Naturale in Verona, Italy, disappeared in the mail. Thus, at 
present, there exists, to the authors' knowledge, no registered material 
of this species. 

Andaniexis pelagica sp.nov. 

(Figs 8-10) 

Andaniexis australis Barnard 1962: 38^40, figs. 24, 25 (immature 9 
& 5.5 mm only, not immature 4.0, 2.0 & 1.5). 



114 



J. BERGE, W. VADER AND A. GALAN 




Fig. 6 Andaniexis gloriosa sp.nov., from: Ledoyer 1986:955, fig. 375 'Andaniexis australis' 



Material EXAMINED. Holotype, Vema Station 53 (see Barnard 
1962). immature female 9.0 mm. Cape Basin, 4893 m. Paratype, 
Vema Station 53, immature 5.5mm. 

DISTRIBUTION. Known from type locality only. 

Dl s( kin ion. Biology: pelagic (Barnard 1962). 

Rostrum very small. 

Antennae elongate. Antenna 1 shorter than antenna 2; flagellum 
3-articulate: accessory Flagellum article 2 absent. Antenna 2 peduncle 



(articles 3-5) longer than flagellum; article 3 short, about as long as 
broad; article 4 shorter than article 5. 

Epistome produced laterally, rectangular with a long ridge on 
each side; epistomal plate (medial keel) produced into a small 
elongate medial ridge exceeding along the entire epistome. 

Mouthparts not elongate or pointed. 

Mandible incisor transverse; incisor smooth; left lacinia mobilis 
present, reduced, laterally straight, not conical. 

Maxilla 1 palp 2-articulate, oval, apex reaching beyond outer 



TYPE MATERIAL OF STEGOCEPHALIDAE 



115 




LMND 



Fig. 7 Andaniexis gloriosa sp.nov., holotype. 



plate; outer plate distally rounded; ST in two parallel rows, first 
marginal and second submarginal, ST first row with 6 setae (ST1-5, 
ST7), ST 1 ordinary (similar to ST 2-4), gap between ST 5 and ST 
7 present, ST A-C present and part of second row; inner plate with 
a weakly developed shoulder, setae pappose. 

Maxilla 2 ordinary, outer plate setae without distal hooks or cleft; 
inner plate setae row A covering the entire margin, appressed to row 
B, row A setae pappose, ?>—4 first setae pectinate distally; row B 
setae proximally pappose, distally with cusps present; row C present; 
row D present, reduced, 1-3 long setae distally, slender. 

Maxilliped palp 4-articulate, article 2 distally unproduced, dacty- 
lus distally simple 

(pointed); inner plate with 2 nodular setae; medial setae-row 
present, not reduced, transverse, simple; distal setae-row present, 
setae simple; inner setae-row present, row reduced to one or two 
setae, conspicuously large and strong; outer plate outer setae-row 
present, marginal, setae attached normally, setae long robust, straight; 
inner setae-row present but strongly reduced, setae short simple, 
slender, appressed to outer setae-row; distal setae-group absent. 

Labrum very short; lobes symmetrical and reduced. 

Labium distally broad, oval. 

Coxal plates and basis of pereopods smooth. Coxae 1-3 
contiguous. 

Pereopod 1 coxa not as deep as basis; propodus subovate. 

Pereopod 2 longer and thinner than pereopod 1 , ischium elongate, 
ratio length:breadth exceeding 1.5, ischium distal posterior margin 
with plumose setae; propodus subrectangular, palm absent. 

Pereopod 4 coxa posteroventral lobe small, reaching about the 
base of the 6th pereon segment; basis anterior and posterior margins 
without long setae, plumose setae on distal anterior and posterior 
margins; ischium with plumose setae on posterior distal margin. 

Pereopod 6 basis posteriorly expanded, expansion rudimentary, 
no row of long plumose setae. 




Fig. 8 Andaniexis pelagica sp.nov., from: Barnard 1962:38, fig. 24 'Andaniexis australis' 



116 



J. BERGE, W. VADER AND A. GALAN 




x E j H 

Fig. 9 Andaniexis pelagica sp.nov., from: Barnard 1962:38, fig. 25 ' Andaniexis australis' 




I iu. 10 Andaniexis pelagica sp.nov.. holotype. 



Pereopod 7 basis anterior margin straight, distally rounded, no 
medial row of setae. 

Oostegites unknown. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus as long as inner. 
Uropod 2 peduncle longer than rami, outer ramus as long as inner. 
Uropod 3 peduncle at least as long as rami, outer ramus 2-articulate, 
outer ramus equal to inner. 

Telson as long as broad, shorter than peduncle uropod 3, entire, 
apically pointed, no submarginal setae apically. 

Male. Unknown. 

REMARKS. Barnard (1962) identified five specimens ('9.0, 5.5, 
4.0, 2.0, and 1.5 mm., sexes unknown' (Barnard 1962: 40) from the 
same Vema station as 'Andaniexis australis (?)', due to the inad- 
equate original description of the latter species (K.H.Barnard 1932). 
However, examination and description of the type material of A. 
australis (see above), together with a closer examination of the 
present material, revealed that it should actually be separated into 
two new species: Andaniexis pelagica (immatures 9.0 and 5.5 mm) 
and Andaniopsis afhcana sp.nov. (see below). [In fact, Barnard 
himself must have reached the same conclusion after he had pub- 
lished his results, as there was a small note in one of the vials 
containing A. pelagica with a note stating 'holotype'.] 

The present species is not a 'typical' member of Andaniexis, i.e. 
it does not belong to the abyssi-gtoup ( Berge & Vader 1 997a), but is 
more closely related to the 'tridentata-gmup' (A. tridentata Ledoyer, 
1986 and A. spinescens Alcock, 1894, see Berge & Vader 1997a), 



TYPE MATERIAL OF STEGOCEPHALIDAE 



117 



and the more distinctive genus Parandaniexis Schellenberg, 1 929. It 
is, however, separated from all Parandaniexis spp by the absence of 
a subchelate pereopod 4. Furthermore, of the above mentioned 
species, only Parandaniexis inermis Ledoyer, 1986 lacks dorsal 
teeth on pleon segments 1-3. 

Andaniopsis Sars, 1895 

Andaniopsis Sars, 1895: 208. 

Andaniopsis Barnard & Karaman 1991: 678. 

Andaniopsis africana sp.nov. 

(Figs 11-12) 

Andaniexis australis Barnard 1962: 38 (part) 

? 'Unknown Genus and Species' Barnard 1967: 150 

MATERIAL EXAMINED. Holotype, immature female 4.0 mm, ' Vema' 
station 53 (see Barnard 1962: 40): Cape Basin, 4893m. Paratypes, 
immature 2.0 and 1.5 mm, 'Vema' station 53. 

DISTRIBUTION. Known only from the type locality (and possibly 
from the west coast of Mexico). Pelagic. 

Description 

Rostrum very small. 

Antennae short. Antenna 1 longer than antenna 2; flagellum 4- 
articulate; accessory flagellum article 2 present. Antenna 2 peduncle 
(articles 3-5) longer than flagellum; article 3 short, about as long as 
broad; article 4 as long as article 5. 



Epistome curved (convex) and smooth; epistomal plate (medial 
keel) produced into a large conspicuous keel. 

Mouthparts not elongate or pointed. 

Mandible incisor transverse; toothed; left lacinia mobilis present, 
powerful, laterally expanded, not conical. 

Maxilla 1 palp 1 -articulate, rectangular, apex not reaching beyond 
outer plate; outer plate distally rounded; ST in two parallel rows, 
first marginal and second submarginal, ST first row with 6 setae 
(ST1-5, ST7), ST 1 ordinary (similar to ST 2-4), gap between ST 5 
and ST 7 present, ST A-C present and part of second row; inner plate 
without a well developed shoulder, setae pappose. 

Maxilla 2 ordinary, outer plate setae without distal hooks or cleft; 
inner plate setae row A covering the entire margin, appressed to row 
B, row A setae pappose, 3-4 first setae weakly pappose proximally, 
slender distally; row B setae proximally simple, distally with cusps 
absent; row C absent; row D absent. 

Maxilliped palp 4-articulate, article 2 distally unproduced, dac- 
tylus distally simple (pointed); inner plate not exceeding base of 
palp article 2; with 2 nodular setae; medial setae-row present, not 
reduced, transverse, setae pectinate; distal setae-row absent; inner 
setae-row present, row reduced to one or two setae, setae con- 
spicuously large and strong; outer plate outer setae-row present, 
marginal, setae attached in a deep hollow, setae short, straight; 
inner setae-row present but strongly reduced, setae short simple, 
slender, parallel but not appressed to outer setae-row; distal setae- 
group absent. 

Labrum not reduced; lobes asymmetrical and reduced. 

Labium distally broad, oval. 




Fig. 11 Andaniopsis africana sp.nov., holotype (see scale 3 on fig. 12). 



18 



J. BERGE, W. VADER AND A. GALAN 



Coxa! plates and basis of pereopods smooth. Coxae 1-3 contigu- 
ous. 

Pereopod 1 coxa deeper than basis: propodus subrectangular. 

Pereopod 2 general appearance like pereopod 1. ischium not 
elongate, ratio length:breadth not exceeding 1.5. ischium distal 
posterior margin with plumose setae; propodus subrectangular, 
palm absent. 

Pereopod 4 coxa posterox entral lobe medium sized, not reaching 
the base of the 7th pereon segment: basis anterior and posterior 
margins without long setae, plumose setae on distal anterior margin, 
no plumose setae on distal posterior margin: ischium without plumose 
setae on posterior distal margin. 

Pereopod 6 basis posteriorly expanded, expansion conspicuous, 
rounded posteriorly, without a row of long plumose setae. 

Pereopod 7 basis anterior margin straight, distally rounded, with 
a medial row of setae, setae short and robust. 

Oostegites on pereopods 2-5; gills on pereopods 2-7. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus as long as inner. 
Uropod 2 peduncle longer than rami, outer 

ramus as long as inner. Uropod 3 peduncle at least as long as rami, 
outer ramus 1 -articulate, outer ramus as long as inner. 

Telson shorter than broad, shorter than peduncle uropod 3, cleft; 
apically rounded, submarginal setae on apex of each lobe. 

Male. Unknown 




Remarks. The present species is described herein in Andaniopsis 
as it resembles Andaniopsis nordlandica in some important 
characters: 1) transverse and toothed incisor and laterally expanded 
lacinia mobilis on the mandible. 2) reduced palp of the first maxilla, 
3) general morphology of the maxilliped, 4) a large epistomal plate 
and 5) relatively long labrum with a well developed right lobe. 
However, the present species possesses a cleft telson, a character not 
found in nordlandica or in the closely related genus Andaniella Sars, 
1895. 

In 1967, Barnard described an immature specimen from a 'un- 
known genus and species' (Barnard 1967: 150). Although the 
epistome and labrum were damaged, and he did not describe the 
telson, the present species seems to be at least closely related to 
Barnard's specimen, if not even belonging to the same species. As 
some essential appendages were damaged, and since it has not been 
possible to examine the specimen, Barnard's specimen is referred to 
as an uncertain synonymy of the present new species. The only 
character that, at present, appears to separate Barnard's specimen as 
a distinct species from africana, is the unexpanded basis on pereopod 
6 (expanded for africana). 

Etymology. The present species is the first species in the genus 
to be recorded from the southern hemisphere and more specifically 
from the coast of South Africa, hence its name africana. 

Bathystegocephalus Schellenberg, 1926 

Bathystegocephalus Schellenberg, 1926: 221 



Fig. 12 Andaniopsis africana sp.nov., holotype: PEP. MX I & MX2: 
stale I.E. EMND. MND&T: scale 2; MXP&AI: scale I on fig, 14. 
P2: scale 3. 




Fig. 13 Bathystegocephalus globosus, female (8mm) syntype. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



119 



Bathystegocephalus globosus (Walker, 1909) 

(Figs 13-15) 

Stegocephalus globosus Walker 1909 
Stegocephaloides valdiviae Strauss 71909 
Bathystegocephalus globosus Schellenberg, 1926 
Bathystegocephalus globosus Pirlot 1933 
Bathystegocephalus globosus Birstein & Vinogradov 1964 

Material examined. Syntypes, BMNH 1909.1.29.17-31 (15 
specimens: females and juveniles, 4.5-10mm) 

Distribution. Indian Ocean. 

Description. Biology: pelagic. 

Rostrum absent. 

Antennae elongate. Antenna 1 shorter than antenna 2: flagellum 
6-articulate; accessory flagellum article 2 absent. Antenna 2 pedun- 
cle (articles 3-5) longer than flagellum; article 3 elongate, article 3 
and 4 geniculate; article 4 shorter than article 5. 

Epistome curved (convex) and smooth; epistomal plate (medial 
keel) produced into a large conspicuous medial keel. 

Mouthparts not elongate or pointed. 

Mandible incisor triangular; incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla 1 palp 1 -articulate, oval, apex reaching beyond outer 
plate; outer plate distally rectangular; ST in two parallel rows, first 
marginal and second submarginal, ST first row with 5 setae, ST 1 
ordinary (similar to ST 2-4), ST 1-5 with one setae absent, ST 6 





Fig. 14 Bathystegocephalus globosus, female (8mm) syntype. 



Fig. 15 Bathystegocephalus globosus, female (8mm) syntype. 

absent, gap between ST 5 and ST 7 present, ST A-D present and part 
of second row; inner plate with a weakly developed shoulder, setae 
pappopectinate. 

Maxilla 2 outer plate absent, inner plate setae row A covering the 
entire margin, appressed to row B, row A setae pappopectinate; row 
B setae proximally pappose, distally with cusps present; row C 
present; row D present, 1-3 long setae distally, with many small 
cusps distally. 

Maxilliped palp 4-articulate, article 2 distally unproduced, dacty- 
lus distally simple (pointed); inner plate without nodular setae; 
medial setae-row present, reduced, transverse, setae pectinate; distal 
setae-row present, reduced, setae simple; inner setae-row present, 
setae conspicuously large, cuspidate; outer plate outer setae-row 
present, marginal, setae attached normally, setae short and straight; 
inner setae-row present, well developed, setae long robust, slender, 
parallel to outer row but widely separated proximally; distal setae- 
group present, setae attached normally, long robust. 

Labrum very short; lobes symmetrical and reduced. 

Labium distally broad, oval. 

Coxal plates and basis of pereopods covered with very short 
setules. Coxae 1-3 contiguous. 

Pereopod 1 coxa deeper than basis; propodus subovate. 

Pereopod 2 general appearance like pereopod 1; ischium not 
elongate, ratio length:breadth not exceeding 1.5, ischium distal 
posterior margin with plumose setae; propodus subovate palm 
absent. 

Pereopod 4 coxa posteroventral lobe medium sized, not reaching 
the base of the 7th pereon segment; basis anterior margin without 
long setae, posterior margin with long setae, without plumose setae 



120 




J. BERGE, W. VADER AND A. GALAN 

Phippsia gibbosa (Sars, 1883) 

(Figs 16-19) 

Stegocephalus gibbosa Sars 1883 

Aspidopleurus gibbosa Sars 1 895 

Material examined. Description based upon immature female, 
Bioice st. 570 (see Berge & Vader 1997d), checked with type (see 
Remarks). 

Distribution. North Atlantic. 

DESCRIPTION. Biology: not pelagic, has been found on living 
Lophelia sp. (see Sars 1895 and W. Vader pers. com.). 

Rostrum very small. 

Antennae short. Antenna 1 longer than antenna 2; flagellum 6- 
articulate; accessory flagellum article 2 absent. Antenna 2 peduncle 
(articles 3-5) longer than flagellum; article 3 short, about as long as 
broad; article 4 shorter than article 5. 

Epistome curved (convex) and smooth; epistomal plate (medial 
keel) produced into a large conspicuous medial keel. 

Mouthparts not elongate or pointed. 

Mandible incisor lateral; incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla 1 palp 2-articulate, rectangular, apex not reaching beyond 
outer plate; outer plate distally rectangular; ST in a pseudocrown 
(see below), ST first row with 6 setae (ST1-5, ST7), ST 1 conspicu- 
ously enlarged, gap between ST 5 and ST 7 absent; ST A and B 
present, located distally, part of first row, ST C present, ST D absent; 
inner plate with a well developed shoulder, setae pappopectinate. 

Maxilla 2 gaping and geniculate, outer plate setae with distal 
hooks present, distal cleft present; inner plate setae row A covering 



Fig. 16 Phippsia gibbosa, immature female, Bioice st.570. 

on distal anterior and posterior margins; ischium with plumose setae 
on posterior distal margin. 

Pereopod 6 elongate, basis posteriorly expanded, expansion rudi- 
mentary, without a row of long plumose setae; carpus and propodus 
anteriorly concave. 

Pereopod 7 basis anterior margin straight, distally rounded, with 
a medial row of long setae. 

Oostegites on pereopods 2-5. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 absent. 
Uropod 1 peduncle shorter than rami, outer ramus longer than inner. 
Uropod 2 peduncle shorter than rami, outer ramus longer than inner. 
Uropod 3 peduncle shorter than half the length of rami, outer ramus 
I -articulate, outer ramus longer than inner. 

Telson as long as broad, as long as peduncle uropod 3, cleft, 
apically rounded, no submarginal setae on apex of each lobe. 

MALE. Pereopod 2 and urosome ordinary (similar to females). 



REMARKS. The present species is distinguishable from all other 
stegocephalid species by its elongate pereopod 6, reduced outer 
plate of maxilla 2 and the triangular shape of the mandibular incisor. 
In addition to this, its long and round coxae I -4 give the anterior part 
of the body a conspicuously globular shape. 

Phippsia Stubbing, 1906 

Aspidopleurm Sars. 1895: 203 (homonym. Pisces) 
Phippsia Stcbbing. 1906: 89 (replacement name) 




MX2 O.P. 



Fig. 17 Phippsia gibbosa, immature female, Bioice st.57(). 



TYPE MATERIAL OF STEGOCEPHALIDAE 



121 




Fig. 18 Phippsia gibbosa. immature female, Bioice st.570. 




Fig. 19 Phippsia gibbosa, immature female, Bioice st.570. 



about two thirds of the margin, clearly separated from row B, row A 
setae pappopectinate; row B setae proximally simple, distally with 
cusps present; row C present; row D present, expanded, row elon- 
gated towards and beyond row A, with many small cusps distally. 

Maxilliped palp 4-articulate, article 2 distal inner margin greatly 
produced, dactylus distally simple (pointed); inner plate with 2 
nodular setae; medial setae-row present, not reduced, vertical, setae 
pectinate; distal setae -row present, setae simple; inner setae-row 
present, row reduced to one or two setae, setae conspicuously large 
and strong; outer plate outer setae-row present, marginal, setae 
attached normally, setae short, strongly curved upwards (hooks); 
inner setae-row present, well developed, setae long robust, setae 
pappose, proximally parallel to outer, distally transverse; distal 
setae-group present, setae attached in a deep hollow, setae short 
simple. 

Labrum elongate; lobes asymmetrical; right lobe not reduced, left 
lobe reduced. 

Labium distally narrowing. 

Coxal plates and basis of pereopods smooth. Coxae 1-3 contigu- 
ous. 

Pereopod 1 coxa deeper than basis; propodus subrectangular. 

Pereopod 2 longer and thinner than pereopod 1; ischium not 
elongate, ratio lengthibreadth not exceeding 1.5, ischium distal 
posterior margin with plumose setae; propodus subrectangular, 
palm absent. 

Pereopod 4 coxa posteroventral lobe very large, reaching beyond 
the base of the 7th pereon segment; basis anterior margin without 
long setae, posterior margin with long setae, without plumose setae 
on distal anterior and posterior margins; ischium with plumose setae 
on posterior distal margin. 

Pereopod 6 basis posteriorly unexpanded, with a row of long 
plumose setae. 

Pereopod 7 basis anterior margin straight, distally rounded, no 
medial row of setae. 

Oostegites on pereopods 2-5. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus longer than inner. 
Uropod 2 peduncle as long as rami, outer ramus shorter than inner. 
Uropod 3 peduncle longer than half the length of rami, outer ramus 
2-articulate, outer ramus shorter than inner. 

Telson longer than broad, longer than peduncle uropod 3, cleft; 
apically rounded, no submarginal setae on apex of each lobe. 

Male. Unknown. 

Remarks. Records at The Natural History Museum, London (see 
also Thurston & Allen, 1969) state that a type specimen of the 
present species is stored there, but that the validity of its status as a 
syntype is doubtful. The specimen in question was collected by 
G.O.Sars from the west coast of Norway, but the absence of a date 
leaves some doubt as to its type status. 

The present species is easily distinguished from all other 
stegocephalid species due to its peculiar gibbous pleonite 3. For a 
revision of this genus, see Berge & Vader (2000). 

Phippsiella Schellenberg, 1925 

Phippsiella Schellenberg 1925: 200 

Phippsiella rostrata K.H. Barnard, 1932 

(Figs 20-21) 

Phippsiella rostrata K.H. Barnard 1932: 76 



122 



J. BERGE, W. VADER AND A. GALAN 




Fig. 20 Phippsiella rostrata, Habitus: female (17mm) paratype. Urosome & L: male 17mm) paralype. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



123 




Fig. 21 Phippsiella roslrata, male (17mm) paratype. 



124 



J. BERGE, W. VADER AND A. GALAN 



Material examined. Syntypes, BMNH 1936.11.2.585-587: 
Discovery St 158 (see K.H. Barnard 1932). 1 male and 2 females 
(17-18 mm). Only the type material is known. 

Distribution. Known only from the type locality (South Georgia). 

Description. Biology: not pelagic, otherwise unknown. 

Rostrum powerful. 

Antennae short. Antenna 1 shorter than antenna 2; flagellum 1 1- 
articulate; accessor) flagellum article 2 present. Antenna 2 peduncle 
(articles 3-5) longer than flagellum: article 3 short.about as long as 
broad: article 4 longer than article 5. 

Epistome curved (convex) and smooth. Epistomal plate (medial 
keel) not produced. Mouthparts not elongate or pointed. 

Mandible incisor lateral: incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla 1 palp 2-articulate, rectangular, apex not reaching beyond 
outer plate: outer plate distally rectangular; ST in two parallel rows, 
first marginal and second submarginal, ST first row with 6 setae 
(ST 1-5, ST7), ST 1 conspicuously enlarged, gap between ST 5 and 
ST 7 present; ST A-C present, part of first row, ST D absent; inner 
plate without a well developed shoulder, setae pappopectinate. 

Maxilla 2 gaping and geniculate, outer plate setae with distal 
hooks present; distal cleft absent; inner plate setae row A covering 
about two thirds of the margin, clearly separated from row B, row A 
setae pappopectinate; row B setae proximally simple, distally with 
cusps present; row C present; row D present, expanded, row elon- 
gated towards and beyond row A, with many small cusps distally. 

Maxilliped palp 4-articulate, article 2 distally unproduced, dacty- 
lus distally simple (pointed); inner plate with 2 nodular setae; medial 
setae-row present, not reduced, transverse, setae pectinate; distal 
setae-row absent; inner setae-row absent; outer plate outer setae- 
row present, marginal, setae attached normally, setae short and 
strongly curved upwards (hooks); inner setae-row present, well 
developed, setae short simple and slender, proximally parallel to 
outer, distally transverse; distal setae-group present, setae attached 
in a deep hollow, setae short simple. 

Labrum elongate; lobes asymmetrical, left lobe weakly reduced. 

Labium distally narrowing. 

Coxal plates and basis of pereopods smooth. Coxae 1-3 contigu- 
ous. 

Pereopod 1 coxa deeper than basis; propodus subrectangular. 

Pereopod 2 general appearance like pereopod 1, ischium not 
elongate, ratio length:breadth not exceeding 1.5, ischium distal 
posterior margin with plumose setae; propodus subrectangular, 
palm absent. 

Pereopod 4 coxa posteroventral lobe medium sized, not reaching 
the base of the 7 ,h pereon segment; basis anterior margin without 
long setae, posterior margin with long setae, plumose setae on distal 
anterior and posterior margins; ischium without plumose setae on 
posterior distal margin. 

Pereopod 6 basis posteriorly expanded, expansion conspicuous, 
rounded posteriorly, with a row of long plumose setae. 

Pereopod 7 basis anterior margin straight, distally rounded, no 
medial row of setae. 

Oostegites on pereopods 2-5. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
1 Iropod I peduncle longer than rami, outer ramus equal to inner. 
I'ropod 2 peduncle longer than rami, outer ramus equal to inner. 
I 'ropod 3 peduncle longer than rami, outer ramus I -articulate, outer 
ramus equal to inner. 

Telson longer than broad, longer than peduncle uropod 3. cleft: 
apically open, no submarginal setae on apex of each lobe. 



Male. Pereopod 2 propodus equally sized in males and females. 
Urosome ordinary (similar to females). 

REMARKS. This species is easily distinguished from all congeners 
by its long rostrum, but it has some affinities with Stegocephalus 
inflatus Kr0yer, 1842. However, the morphology of both the 
mouthparts and of the basis of pereopod 7 strongly suggests that this 
species should be retained in the genus Phippsiella. 

Stegocephaloides Sars, 1895 

Stegocephaloides Sars 1895: 201 

Stegocephaloides attingens K.H. Barnard, 1932 

(Figs 22-25) 

Stegocephaloides attingens K.H. Barnard 1932: 131 (3 of 4 speci- 
mens only) 
? Stegocephaloides attingens Barnard 1961: 60 
? Stegocephaloides attingens Griffiths 1975: 167 

MATERIAL EXAMINED. Syntypes, BMNH 1928.4.13.41-44; 3 
specimens (but see below): female, male and immature, 5-8 mm, 
Cape Point, South Africa, 500- 1000m. Additional material: SAM 
A4423, 4 specimens, Cape Point. 

Distribution. South Africa to (?) Angola, 990-1 400m. 




Fig. 22 Stegocephalides attingens, female (8mm) cotype. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



125 




Fig. 23 Stegocephalides attingens, female (8mm) cotype. 



DESCRIPTION. Biology: not pelagic, otherwise unknown. Based 
on cotype, female 8 mm. 

Rostrum very small. 

Antennae short. Antenna 1 equal to antenna 2; flagellum 4- 
articulate; accessory flagellum article 2 absent. Antenna 2 peduncle 
(articles 3-5) longer than flagellum; article 3 short, about as long as 
broad; article 4 about as long as 5. 



Epistome curved (convex) and smooth; epistomal plate (medial 
keel) not produced. Mouthparts not elongate or pointed. 

Mandible incisor lateral; incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla 1 palp 1 -articulate, rectangular, apex not reaching beyond 
outer plate and apex reaching beyond outer plate; outer plate distally 
rectangular; ST in a pseudocrown, ST first row with 6 setae (ST1-5, 



126 



J. BERGE, W. VADER AND A. GALAN 




Fig. 24 Stegocephalid.es attingens, female (8mm) cotype. 

ST7), ST 1 conspicuously enlarged, gap between ST 5 and ST 7 
present: ST A present, located distally. part of first row; ST B-C 
present, part of second row. ST D absent; inner plate without a well 
developed shoulder, setae pappose. 

Maxilla 2 gaping and geniculate, outer plate setae with distal 
hooks present, distal cleft absent; inner plate setae row A covering 
the entire margin, clearly separated from row B, row A setae 
pappopectinate; row B setae proximally pappose, distally with 
cusps present; row C present; row D present, 1-3 long setae distally, 
setae with many small cusps distally. 

Maxilliped palp 4-articuIate, article 2 distal inner margin weakly 
produced, dactylus distally simple (pointed); inner plate with 2 
nodular: medial setae-row present, reduced to one or two setae but 
differentiated from distal row, transverse, setae pectinate; distal 
setae-row present, setae simple; inner setae-row present, setae not 
conspicuously large; outer plate outer setae-row present, marginal, 
setae attached in a deep hollow, setae short, strongly curved upwards 
(hooks); inner setae-row present but strongly reduced, setae short 
simple and slender, proximally parallel to outer, distally transverse; 
distal setae-group present, setae attached in a deep hollow, setae 
short simple. 

Labium elongate: lobes asymmetrical, right lobe not reduced, left 
lobe reduced. 

Labium distally narrowing. 

Coxal plates and basis ol pereopods covered with simple setae. 
Coxae I -3 contiguous. 



Fig. 25 Stegocephalides attingens, female (8mm) cotype. 



Pereopod 1 coxa deeper than basis; propodus subovate. 

Pereopod 2 general appearance like pereopod 1; ischium not 
elongate, ratio length:breadth not exceeding 1.5; ischium distal 
posterior margin plumose setae present; propodus subrectangular; 
palm absent. 

Pereopod 4 coxa posteroventral lobe large, reaching about the 
base of the 7th pereon segment; basis anterior margin without long 
setae, posterior margin with long setae, plumose setae on distal 
anterior margin, no plumose setae on distal posterior margin; is- 
chium with plumose setae on posterior distal margin. 

Pereopod 6 basis posteriorly unexpanded, with a row of long 
plumose setae present. 

Pereopod 7 basis anterior margin straight, distally pointed and 
acute, with medial row of long setae. 

Oostegites on pereopods 2-5, but strongly reduced on pereopod 
2. Gills on pereopods 2-7. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 absent. 
Uropod 1 peduncle longer than rami, outer ramus as long as inner. 
Uropod 2 peduncle longer than rami, outer ramus shorter than inner. 
Uropod 3 peduncle longer than half the length of rami, outer ramus 
1 -articulate, outer ramus longer than inner. 

Telson longer than broad, longer than peduncle uropod 3, cleft, 
apically pointed, submarginal setae on apex of each lobe. 

Male. Pereopod 2 propodus larger in males than in females. 
Urosome ordinary (similar to females). 

Remarks. The distally acute basis on pereopod 7 is a character 
state only shared with Stegocephaloides auratus (Sars, 1 883), but S. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



127 



attingens is separated from the latter species by the shape of coxa 4 
(S. auratus has a very deep coxa with the lower margin straight, 
whereas in 5. attingens coxa 4 is not as deep and the lower margin is 
curved), by the strongly reduced oostegite on pereopod 2, and by 
article 4 on antenna 2 being almost as long as article 5. 

Stegocephaloides australis K.H. Barnard, 1916 

(Figs 26-28) 



Stegocephaloides australis K.H. Barnard 1916: 129 
? Stegocephaloides australis Griffiths 1974: 324 
? Stegocephaloides australis Griffiths 1975: 167 
Non Stegocephaloides australis Ledoyer 1986: 962 

Material examined. Syntypes, BMNH 1928.4.13.45^48; 4 fe- 
males (6-8 mm), Cape Point, South Africa, 110-200m. 

Distribution. Endemic to South Africa. 




/ y. 




Fig. 26 Stegocephaloides australis, female (7mm) cotype. 



128 



J. BERGE, W. VADER AND A. GALAN 




Fig. 27 Stegocephaloid.es australis, female (7mm) cotype. 

Description. Biology: not pelagic, otherwise unknown. 

Rostrum very small. 

Antennae short. Antenna 1 equal to antenna 2; flagellum 5- 
articulate; accessory 

flagellum article 2 absent. Antenna 2 peduncle (articles 3-5) 
longer than flagellum; article 3 short, about as long as broad; article 
4 as long as article 5. 

Epistome curved (convex) and smooth; epistomal plate (medial 
keel) not produced. 

Mouthparts not elongate or pointed. 

Mandible incisor lateral; incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla I palp 1 -articulate, rectangular, apex not reaching beyond 
outer plate; outer plate distally rectangular; ST in a pseudocrown, 
ST first row with 6 setae (ST1-5, ST7), ST 1 conspicuously en- 
larged, gap between ST 5 and ST 7 present; ST A present, located 
distally and part of first row, ST B present, part of second row, STC 
present. ST D absent; inner plate with a weakly developed shoulder, 
setae pappopectinate. 

Maxilla 2 gaping and geniculate, outer plate setae with distal 
hooks present, distal cleft absent; inner plate setae row A covering 
about two thirds of the margin, clearly separated from row B, row A 
setae pappopectinate; row B setae proximally pappose, distally with 
cusps present; row C present; row D present, expanded, row elon- 
gated towards and beyond row A, with many small cusps distally. 

Maxilliped palp 4-articulate, article 2 distal inner margin weakly 
produced, dactylus distally cleft with one pointed and one heavily 
setose part; inner plate with 2 nodular setae; medial setae-row 



present, reduced to one or two setae but differentiated from distal 
row, transverse, setae pectinate; distal setae-row present, setae 
simple; inner setae-row present, setae not conspicuously large; outer 
plate outer setae-row present, marginal, setae attached in a deep 
hollow, setae short, strongly curved upwards (hooks); inner setae- 
row present but strongly reduced, setae short simple and slender, 
proximally parallel to outer row and distally transverse; distal setae- 
group present, setae attached in a deep hollow, setae short simple. 

Labrum elongate; lobes asymmetrical, right lobe not reduced, left 
lobe reduced. 

Labium distally narrowing. 

Coxal plates and basis of pereopods smooth. Coxae 1-3 contigu- 
ous. 

Pereopod 1 coxa deeper than basis; propodus subrectangular. 

Pereopod 2 general appearance like pereopod 1, ischium not 
elongate, ratio 

length:breadth not exceeding 1 .5, ischium distal posterior margin 
with plumose setae; propodus subrectangular, palm absent. 

Pereopod 4 coxa posteroventral lobe large, reaching about the 
base of the 7th pereon segment; basis anterior margin without long 
setae, posterior margin with long setae, plumose setae on distal 
anterior and posterior margins; ischium with plumose setae on 
posterior distal margin present. 

Pereopod 6 basis posteriorly expanded, expansion rudimentary, 
with a row of long plumose setae. 

Pereopod 7 basis anterior margin straight, distally rounded, with 
a medial row of short and robust setae. 




Fig. 28 Stegocephaloides australis, female (7mm) cotype. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



129 



Oostegites on pereopods 2-5. Gills on pereopods 2-7. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 absent. 
Uropod 1 peduncle longer than rami, outer ramus as long as inner. 
Uropod 2 peduncle longer than rami, outer ramus as long as inner. 
Uropod 3 peduncle longer than half the length of rami, outer ramus 
2-articulate, outer ramus as long as inner. 

Telson longer than broad, longer than peduncle uropod 3, cleft, 
apically pointed, no submarginal setae on apex of each lobe. 

Male. Unknown. 



Remarks. This species is distinguished from all other stego- 
cephalid species by the very peculiar dactylus on its maxilliped palp 
(see figure 27). 

Stegoceph.aloid.es calypsonis sp. nov. 

(Figs 29-30) 

Material examined. Holotype, female, 4 mm, BMNH 
1992:1483:17, 10°38'N 65°32'W, 1400m, Calypso exp 7 th . October 
1979. Paratypes, 18 specimens, BMNH 1992:1483:17 (details, as 
above). Additional material: BMNH 1992:1484:15, 11 specimens, 
10°31'N 64°45'W, 1300m, Calypso exp. 5 lh October 1979 

DISTRIBUTION. Known only from the type locality. 

DESCRIPTION. Biology: not pelagic, otherwise unknown. 

Rostrum very small. 

Antennae elongate. Antenna 1 longer than antenna 2; flagellum 6- 
articulate; accessory flagellum article 2 present. Antenna 2 peduncle 
(articles 3-5) longer than flagellum; article 3 

short, about as long as broad; article 4 shorter than article 5. 

Epistome curved (convex) and smooth; epistomal plate (medial 
keel) not produced. 

Mouthparts not elongate or pointed. 

Mandible incisor lateral; incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla 1 palp 1 -articulate, rectangular, apex not reaching beyond 
outer plate; outer plate distally rectangular; ST in a pseudocrown, 
ST first row with more than 6 setae (ST1-5 expanded, ST7), ST 1 
conspicuously enlarged, gap between ST 5 and ST 7 absent; ST A 
present, located distally and part of first row, ST B present, part of 
second row, ST C present, ST D absent; inner plate without or with 
a weakly developed shoulder, setae pappose. 

Maxilla 2 gaping and geniculate, outer plate setae with distal 
hooks present, distal cleft absent; inner plate setae row A cover- 
ing about two thirds of the margin, clearly separated from row B, 
row A setae pappopectinate; row B setae proximally pappose, 
distally with cusps present; row C present; row D present, ex- 
panded, row elongated towards and beyond row A, with many 
small cusps distally. 

Maxilliped palp 4-articulate, article 2 distally unproduced, dacty- 
lus distally simple (pointed); inner plate with 1 nodular setae; medial 
setae-row present, reduced to one or two setae but differentiated 
from distal row, transverse, setae pectinate; distal setae-row present, 
setae simple; inner setae-row present, setae not conspicuously large; 
outer plate outer setae-row present, marginal, setae attached in a 
deep hollow, setae short and straight; inner setae-row present but 
strongly reduced, setae short and simple, proximally parallel to 
outer and distally transverse; distal setae-group present, setae at- 
tached in a deep hollow, setae short simple. 

Labrum elongate; lobes asymmetrical; right lobe not reduced; left 
lobe reduced. 

Labium distally narrowing. 



Coxal plates and basis of pereopods smooth. Coxae 1-3 con- 
tiguous. 

Pereopod 1 coxa deeper than basis; propodus subovate. 

Pereopod 2 longer and thinner than pereopod 1 ; ischium elongate, 
ratio length: breadth exceeding 1.5, ischium distal posterior margin 
with plumose setae; propodus subrectangular, palm absent. 

Pereopod 4 coxa posteroventral lobe medium sized, not reaching 
the base of the 7 lh pereon segment; basis anterior margin without 
long setae, posterior margin with long setae present, no plumose 
setae on distal anterior and posterior margins; ischium without 
plumose setae on posterior distal margin. 

Pereopod 6 basis posteriorly unexpanded, without a row of long 
plumose setae. 

Pereopod 7 basis anterior margin straight, distally rounded, no 
medial row of setae. 

Oostegites on pereopods 2-5. Gills on pereopods 2-7. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 absent. 
Uropod 1 peduncle longer than rami, outer ramus shorter than inner. 
Uropod 2 peduncle shorter than rami, outer ramus shorter than inner. 
Uropod 3 peduncle longer than half the length of rami, outer ramus 
2-articulate, outer ramus shorter than inner. 

Telson longer than broad, as long as peduncle uropod 3, cleft, 
apically rounded, no submarginal setae on apex of each lobe. 

MALE. Pereopod 2 propodus larger in males than in females. 
Urosome ordinary (similar to females). 

Remarks. Stegocephaloides calypsonis is a rather small species, 
but with somewhat elongate appendages (antennae and pereopods). 
The arrangement of ST on maxilla 1 is similar to that found in S. 
christianiensis (Boeck, 1 87 1 ), but the presence of a second article on 
outer ramus of uropod 3 and the rounded lower margin of coxa 4 
indicate some relationship with S. attingens (outer ramus articula- 
tion absent, but with rounded coxa), S. boxshalli (see below) and S. 
camoti (Barnard, 1967). 

Etymology. Named after the French oceanographic vessel 
'Calypso' from which the material was sampled. Galan (in his 
unpublished 1984 Ph.D. thesis) originally used the name calypsae, 
but this is here emended to the grammatically more correct 
calypsonis. 

Stegocephaloides boxshalli sp. nov. 

(Figs 31-33) 

Stegocephaloides attingens K.H. Barnard 1932: 131 (1 out of 4 
specimens only) 

Material examined. Holotype, SAM A43988, male, Cape Point 
20 th of August 1903. 

Paratypes, SAM A43988, 3 specimens. Additional material: 
BMNH 1928.4.13.41^14, female ~7mm: Cape Point, South Africa, 
500- 1000m. 

Distribution. Known only from Cape Point, South Africa. 

Description. Biology: not pelagic, otherwise unknown. 

Rostrum very small. 

Antennae short. Antenna 1 longer than antenna 2; flagellum 5- 
articulate; accessory flagellum, article 2 present. Antenna 2 peduncle 
(articles 3-5) as long as flagellum; article 3 short, about as long as 
broad; article 4 longer than article 5. 

Epistome curved (convex) and smooth; epistomal plate (medial 
keel) not produced. 

Mouthparts not elongate or pointed. 



130 



J. BERGE. W. VADER AND A. GALAN 




Fig. 2 f > Sh ■••' ephaloides calypsonis sp.nov., holoiype. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



131 




Fig. 30 Stegocephaloides calypsonis sp.nov., holotype. 

Mandible incisor lateral; incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla 1 palp 1 -articulate, rectangular, apex not reaching beyond 
outer plate; outer plate distally rectangular; ST in two parallel rows, 
first marginal and second submarginal, ST first row with 7 setae 
(ST1-5, ST7 and an additional ST located medially on outer plate), 
ST 1 ordinary (similar to ST 2-4), gap between ST 5 and ST 7 
present; ST A-C, part of second row. ST D absent; inner plate with 
a well developed shoulder, setae pappopectinate. 

Maxilla 2 gaping and geniculate, outer plate setae with distal 
hooks present, distal cleft absent; inner plate setae row A covering 
about two thirds of the margin, clearly separated from row B, row A 
setae pappopectinate; row B setae proximally simple, distally with 
cusps present; row C present; row D present, expanded and elon- 
gated beyond row A, with many small cusps distally. 

Maxilliped palp 4-articulate, article 2 distal inner margin weakly 
produced, dactylus distally simple (pointed); inner plate with 2 
nodular setae; medial setae-row absent, distal setae-row present, 
setae simple; inner setae-row present, row reduced to one or two 
setae, setae not conspicuously large; outer plate outer setae-row 
present, marginal, setae attached in a deep hollow, setae short and 
strongly curved upwards (hooks); inner setae-row present but strongly 
reduced, setae short simple and slender, proximally parallel to outer 
and distally transverse; distal setae-group present, setae attached in 
a deep hollow, setae short simple. 

Labrum elongate; lobes asymmetrical; right lobe not reduced; left 
lobe reduced. 

Labium distally narrowing. 

Coxal plates and basis of pereopods covered with very short 
setules. Coxae 1-3 contiguous. 



Pereopod 1 coxa deeper than basis; propodus subovate. 

Pereopod 2 longer and thinner than pereopod 1, ischium not 
elongate, ratio length:breadth not exceeding 1.5, ischium distal 
posterior margin with plumose setae; propodus subovate, palm 
absent. 

Pereopod 4 coxa posteroventral lobe large, reaching about the 
base of the 7 lh pereon segment; basis anterior margin without long 
setae, posterior margin with long setae, no plumose setae on distal 
anterior and posterior margins; ischium with plumose setae on 
posterior distal margin. 

Pereopod 6 basis posteriorly expanded, expansion rudimentary, 
with a row of long plumose setae. 

Pereopod 7 basis anterior margin straight, distally rounded, with 
a medial row of short and robust setae. 

Oostegites on pereopods 2-5. Gills on pereopods 2-7. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus longer than inner. 
Uropod 2 unknown. Uropod 3 peduncle longer than rami, outer 
ramus 2-articulate, outer ramus shorter than inner. 

Telson as long as broad, longer than peduncle uropod 3, cleft, 
apically rounded, no submarginal setae on apex of each lobe. 

Male. Pereopod 2 propodus larger in males than in females. 
Urosome ordinary (similar to females). 

REMARKS. All five specimens were unfortunately lost in the mail 
after examination of the material, but before the description of the 
species was entirely finished. Four slides made from the holotype 
represent the only remaining material. Thus figures of uropod 2, 
epimeral plate 1 & 2 and habitus are not available. 

The present species has some affinities to S. attingens (see above), 
reflected in the fact that all five specimens registered for this species 
had initially been identified as S. attingens (indeed, the one speci- 
men identified by K.H. Barnard (1932) was among the type material 
of S. attingens). The two species are, however, separated by the basis 
of pereopod 7 (S. boxshalli is not pointed distally), by uropod 3 {S. 
boxshalli has a 2-articulate outer ramus) and by the shorter and more 
rounded telson of S. boxshalli. Furthermore, S. boxshalli can be 
separated from all other stegocephalid species by the presence of an 
additional ST on the outer margin of maxilla 1 outer plate. 

Etymology. The present species is named after Prof. Geoff 
Boxshall (Natural History Museum in London) for his support and 
help with the first authors' work on his Ph.D. thesis. 

Stegocephaloides ledoyeri sp. no v. 

(Figs 34-36) 

Stegocephaloides australis Ledoyer 1986:962 (non S. australis 
K.H. Barnard) 

Material examined. Holotype, SAM A 15598, female 8mm. 
27°59.5'S 32°40.8'E, collected 22 nd May 1976. Additional material: 
14 specimens identified as S. australis, see Ledoyer 1986:962. 

Distribution. Known from South Africa and Madagascar. 

DESCRIPTION. Biology: not pelagic, otherwise unknown. 

Rostrum very small. 

Antennae short. Antenna 1 shorter than antenna 2 and longer than 
antenna 2; flagellum 6-articulate; accessory flagellum article 2 
present. Antenna 2 peduncle (articles 3-5) longer than flagellum; 
article 3 short, about as long as broad; article 4 shorter than article 5. 

Epistome curved (convex) and smooth; epistomal plate (medial 
keel) not produced. 



132 



J. BERGE, W. VADER AND A. GALAN 




Fin. 31 Sh -"• ephaloides boxshalli sp.nov.. holotypc. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



133 





Fig. 32 Stegocephaloides boxshalli sp.nov., holotype. 



Fig. 33 Stegocephaloides boxshalli sp.nov., holotype. 



Mouthparts not elongate or pointed. 

Mandible incisor lateral; incisor toothed; left lacinia mobilis 
present, powerful, laterally expanded, not conical. 

Maxilla 1 palp 1 -articulate, rectangular, apex not reaching beyond 
outer plate; outer plate distally rectangular; ST in a pseudocrown, 
ST first row with 6 setae (ST1-5, ST7), ST 1 conspicuously en- 
larged; ST A present; located distally and part of first row, ST B-C 
present, part of second row, ST D absent; inner plate with a weakly 
developed shoulder, setae pappopectinate. 

Maxilla 2 gaping and geniculate, outer plate setae with distal 
hooks present, distal cleft absent; inner plate setae row A covering 
about two thirds of the margin, clearly separated from row B, row A 
setae pappopectinate; row B setae proximally pappose, distally with 
cusps present; row C present; row D present, expanded and elon- 
gated beyond row A, with many small cusps distally. 

Maxilliped palp 4-articulate, article 2 distally produced, distal 
inner margin weakly produced, dactylus distally simple (pointed); 
inner plate with 2 nodular setae; medial setae-row present, re- 
duced to one or two setae but differentiated from distal row, 
transverse, setae pectinate; distal setae-row present, setae simple; 
inner setae-row present, setae not conspicuously large; outer plate 
outer setae-row present, marginal, setae attached in a deep hollow, 
setae short and strongly curved upwards (hooks); inner setae-row 
present but strongly reduced, setae short, simple and slender, 
proximally parallel to outer and distally transverse; distal setae- 



group present, setae attached in a deep hollow, setae short simple. 

Labrum elongate; lobes asymmetrical, right lobe not reduced, left 
lobe reduced. 

Labium distally narrowing. 

Coxal plates and basis of pereopods smooth. Coxae 1-3 contigu- 
ous. 

Pereopod 1 coxa deeper than basis; propodus subovate. 

Pereopod 2 longer and thinner than pereopod 1, ischium not 
elongate, ratio length:breadth not exceeding 1.5, ischium distal 
posterior margin with plumose setae; propodus subrectangular, 
palm absent. 

Pereopod 4 coxa postero ventral lobe large, reaching about the 
base of the 7th pereon segment; basis anterior margin without long 
setae, posterior margin with long setae, plumose setae on distal 
anterior margin, no plumose setae on distal posterior margin; is- 
chium with plumose setae on posterior distal margin. 

Pereopod 6 basis posteriorly expanded, expansion rudimentary, 
with a row of long plumose setae present. 

Pereopod 7 basis anterior margin straight, distally rounded, with 
a medial row of short and robust setae. 

Oostegites on pereopods 2-5. Gills on pereopods 2-7. 

Pleonites 1-3 dorsally smooth. 

Urosome: articulation between urosome segments 2 and 3 present. 
Uropod 1 peduncle longer than rami, outer ramus as long as inner. 
Uropod 2 peduncle as long as rami, outer ramus shorter than inner. 



134 



J. BERGE, W. VADER AND A. GALAN 




He. 34 Si, ephaloides ledoyeri sp.nov., from: Ledoyer 19X6:963. fig. 379 'Stegocephaloides australis'. 



TYPE MATERIAL OF STEGOCEPHALIDAE 



135 





Fig. 35 Stegocephaloides ledoyeri sp.nov., holotype. 



Fig. 36 Stegocephaloides ledoyeri sp.nov., holotype. 



Uropod 3 peduncle longer than half the length of rami, outer 
ramus 1 -articulate, outer ramus longer than inner. 

Telson longer than broad, longer than peduncle uropod 3, cleft, 
apically pointed, no submarginal setae on apex of each lobe. 

Male. Pereopod 2 propodus larger in males than in females. 
Urosome ordinary (similar to females). 

Remarks. The present species is distinguished from all other 
Stegocephaloides species by the combination of a distally rounded 
basis of pereopod 7 and a short article 4 on the peduncle of the 
second antenna (about 1/2 the length of article 5). 

Etymology. This species is named after Prof. M. Ledoyer, who 
first described material of this species (Ledoyer 1 986:962, identified 
as Stegocephaloides attingens). 



DISCUSSION 

The present paper is part of series that will lead to a complete 
revision of the family (Berge & Vader, in press.), that will also 
include a cladistic analysis of all its species. Thus, the species treated 
herein are described without any further reference to their 
phylogenetic relationships. Consequently, some of these species 
may, in future, be transferred to different genera. 



REFERENCES 



Barnard, J. L. 1961. Gammaridean Amphipoda from depths of 400 to 6000 meters. 

Galathea Report 5 :23-128. 
. 1962. South Atlantic abyssal amphipods collected by R. V. Vema. Vema Research 

Series 1 : 1-78. 
. 1964. Deep-sea Amphipoda (Crustacea) collected by the R/V «Vema» in the 

eastern Pacific Ocean and the Caribbean and Mediterranean seas. Bulletin of the 

American Museum of Natural History 127 ( 1 >: 1 — 46. 
. 1967. Bathyal and abyssal gammaridean Amphipoda of Cedros Trench, Baja 

California. United States National Museum Bulletin 260 : 1-205. 
& G. S. Karaman. 1991. The families and genera of marine gammaridean 

Amphipoda (except marine gammaroids). Part 2. Records of the Australian Museum 

Supplement 13 (2):419-866. 
Barnard, K. H. 1916. Contributions to the crustacean fauna of South Africa. Annals of 

the South African Museum 15 (3): 105-302. 

. 1932. Amphipoda. Discovery Reports 5 : 1-326. 

Berge, J. 2001a. Revision of the Amphipod (Crustacea: Stegocephalidae) genera 

Andaniotes Stebbing, 1897 and Metandania Stephensen, 1925. Journal of Natural 

History 35 :787-832. 
. 2001b. Revision of Stegosoladidus Barnard & Karaman, 1987 (Crustacea: 

Amphipoda: Stegocephalidae); Redescription of two species and description of 

three new species. Journal of Natural History 35 :539— 571. 
& Vader, W. 1997a. Atlantic and Mediterranean species of the genus Andaniexis 

Stebbing (Amphipoda: Stegocephalidae). Journal of Natural History 31 :1429- 

1455. 
. 1997b. North Atlantic and Mediterranean species of the genus Phippsielia 

Schellenberg (Amphipoda: Stegocephalidae). Journal of Natural History 31 : 1501- 

1532. 
. 1997c. North Atlantic species of the genus Stegocephaloides Sars (Amphipoda. 

Stegocephalidae). Sarsia 82 :325-346. 
. 1997d. Stegocephalid (Crustacea, Amphipoda) species collected in the BIOFAR 



136 J. BERGE, W. VADER AND A. GALAN 

and BIOICE programmes. Sarsia 82 :347-370. Caprellidca of (he Cape Province east of Cape Agulhas. Annuls of the South African 
. 2(HX) Revision of the Stegocephalid (Crustacea: Amphipoda) genera Phippsia Museum 65 (9):251-336. 

and IctraJeion. with descriptions of 4 new species Memoirs of the Museum of . 1975. The Amphipoda of southern Africa. Part 5. The Gamniaridea and Caprellidea 

\ ictoria 58 : 149-178. of the Cape Province west of Cape Agulhas. Annals of the South African Museum 67 
. in press Revision of the Amphipod (Crustacea) family Stegocephalidae. Zoo- (5):91— 181. 

logical Journal oj the l.innncan Society. Ledoyer, M. 1986. Crustaces Amphipodes Gammariens. Families des Haustoriidae a 

Griffiths. C. L. 1974. The Amphipoda of southern Africa. Part 4. The Gamniaridea and Viljazianidae. Faune <le Madagascar 59 (2 1:599-1 112. 



Bull. nat. Hist. Mus. Lond. (Zool.) 67(2): 137-K 



^(2^cS2. \ ^ 



Issued 29 November 2001 



The genus Ischioscia Verhoeff, 1928 in 
Venezuela, with the description of six new 
species (Crustacea, Oniscidea, Philosciidae) 

ANDREAS LEISTIKOW 

Universitdt Bielefeld, Abteilung fur Zoomorphologie und Systematik, Morgenbreede 45, D-33615 Bielefeld, 
Germany and Ruhr-Universitat Bochum, Lehrstuhl fiir Spezielle Zoologie, Universitdtsstrafie 150, D-44780 
Bochum, Germany, e-mail: Leiste@Biologie.Uni-Bielefeld.de 

CONTENTS 

Introduction 137 

Materials and Methods 137 

Systematics 138 

Ischioscia variegata (Dollfus, 1893) 138 

Ischioscia fasciifrons sp. nov 138 

Ischioscia hirsuta sp. nov 143 

Ischioscia colorata sp. nov 143 

Ischioscia pariae sp. nov 152 

Ischioscia guamae sp. nov 157 

Ischioscia trifasciata sp. nov 157 

Discussion 166 

Acknowledgements 167 

References 167 



SYNOPSIS. Terrestrial isopods (Oniscidea) collected in Venezuela in 1998 revealed plentiful material of the philosciid genus 
Ischioscia. This genus was, until very recently, represented in Venezuela by only a single species. /. variegata (Dollfus, 1893). 
The examination of the new samples lead to the description of six new species, all confined to small areas within Venezuela; five 
of which are closely related to /. variegata. These species are described in the present paper and the biogeography of the genus 
is discussed for Venezuela. The characters, which are important for reconstructing the phylogeny are presented and the 
relationships of the species are also discussed. 



INTRODUCTION 



The genus Ischioscia Verhoeff, 1928 was one of the first genera of 
philosciid Oniscidea described from South America (Verhoeff 1928). 
It was instituted for a species from Venezuela, /. lobifera Verhoeff, 
1928, which is now considered to be a junior synonym of /. 
variegata (Dollfus, 1893), a species reported from several localities 
in northern South America (Leistikow 1997). The genus comprises 
the largest species of terrestrial Isopoda in South America, with /. 
variegata reaching a body length of about 15 mm. Beside this 
species, several others are reported from the Peruvian and Brazilian 
Amazon region (Lemos de Castro 1955, Schmalfuss 1980), from 
Central America (van Name 1926, Arcangeli 1930, Leistikow 1997, 
1999 and 2000) and even from the lesser Antilles (van Name 1936). 
The total number of nominal species now is 16, and it is likely that 
several others will be found when the vast Amazonas region is better 
explored. The number of known species increased in those regions 
where larger collections were made. For example, in Costa Rica, ten 
species have been discovered in the last decade. The samples from 
Venezuela dealt with in this study revealed the presence of six more 
species; most of them close to /. variegata, but quite distinct in 
several characters. Hence, several records of/, variegata may show 



to belong to other species. The species of Ischioscia found in 
Venezuela are described in detail, the holotypes are deposited in The 
Natural History Museum, London for which the acronym BMNH is 
used below. Paratypes are deposited in The Natural History Mu- 
seum, London, the Museum d'Histoire Naturelle, Geneve (MHNG), 
Staatliches Museum fur Naturkunde, Stuttgart (SMNS), the mu- 
seum of the University of Marcay, Venezuela (MUMV), and in the 
collection of the author. The new species belong to the martinae- 
variegata-group of species distributed in Central America and 
northern South America. Particularly five of the new species are 
forming a monophylum together with /. variegata and /. panamensis. 
They seem to be restricted to smaller areas in Venezuela, and their 
distribution is compared with those of other taxa. 



MATERIALS AND METHODS 

Several samples of isopods preserved in 70% ethanol were examined. 
The samples were checked with a Wild stereoscope and in the case of 
new species a holotype was selected for preparation. Drawings were 
made using a camera lucida. Some specimens were dissected and the 
appendages were mounted on microscopic slides in glycerine gelatine. 



© The Natural History Museum, 2001 



138 



A. LEISTIKOW 



the appendages wore then drawn using a camera lucida including 
draw nigs of the new species. The types selected for museum storage 
are preserved in 70% ethanol/5% glycerine. 



SYSTEMATICS 

Ischioscia variegata (Dollfus, 1893) 

Material EXAMINED. 19 males, 1 1 females (with marsupium), 1 1 
females/immatures: Parque Nacional El Avila 10°34.70'N 
66°53.92'W, banana plantation, forest along brook, leg. C. Schmidt 
10.3.1998: 3 males. 10 females (with marsupium). (MHNG); 8 
females 1 1 miniatures: 10 o 33.01'N66 o 54.41'W Forest with Bamboo, 
Heliconia sp., Ricinus sp. among others, gas pipeline crossing under 
street, small river, under bamboo leaves on sand, gravel, leg. C. 
Schmidt 1 1.3.1998; 1 male: Aragua, Parque Nacional Henry Pittier. 
Mist forest near Estacion Biologica Rancho Grande, between leaves 
of fallen bromeilad. leg. C. Schmidt 14.3.1998; 1 male, 1 females 
(with marsupium). 1 immature: Aragua, Parque Nacional Henry 
Pittier, street heading from Rancho Grande to sea shore, nearCapilla 
Virgen del Carmen. Mist forest, brook with cascade, under moist leaf 
litter and stones, leg. C. Schmidt 15.3.1998; 1 male: Falcon, Parque 
Morrocoy, Peninsula de Morrocoy, northern shore, Cueva del Indio 
i carstic cave with crushed ceiling, within detritus in small niches and 
edges of the rock, under stones on bottom, leg. C.Schmidt 18.03.1998; 
3 males, 3 females (with marsupium), 6 females: Sierra de San Luis, 
11°07.35'N 69°40.74'W opposite to Cueva San Luis, forest with 
almost dried out brook emerging from cave, stones litter, 
Bombacaceae, Piperaceae, mango-trees, under logg, leg. C. Schmidt 
20.3.1998: 2 males, 1 females (with marsupium), 2 females: 
Curimagua 11°1 1.61'N 69°39.96'W track to Cueva de Lugo, banana 
plantation, under loggs, leg. C. Schmidt 21.3.1998; 1 female, 3 
immatures: Curimagua, Cueva de Lugo 11°11.90'N 69°39.88'W 
carstic cave without water, moist bottom, at the entrance under 
stones, leg. C. Schmidt 21.3.1998; 4 males, 3 females (with marsu- 
pium), 6 females, 2 immatures: Cueva Acurite 1 1 ° 10.42'N 69°37.75'W 
cartic cave with water, surrounded by forest and banana plantation, 
at the entrance under stones and leaf litter, leg. C. Schmidt 21.3.1 998; 
3 males, 3 females, 1 immature: Eastern slopeof Andes 'La Campana' 
8°51.92'N 70°37.14'W 1500 m (+/- 200 m) moist forest (Araceae, 
Arecaceae, Melostomataceae, Orchidaceae) under leaf litter, leg. C. 
Schmidt 24.3.1998; 1 male, 1 female: Penfnsula de Paria, Puy Puy 
10°42.00'N 62°58.05'W bay with sandy beach, coconut palms, 
surrounded by dry mountains, small westernmost bay, under loggs, 
coconuts and leaf litter, leg. C. Schmidt 29.3. 1 998; 1 males, 3 females 
(with marsupium), 1 1 females/immatures: Peninsula de Paria, Puy 
Puy 10°42.00'N 62°58.05'W bay with sandy beach, coconut palms, 
surrounded by dry mountains, forest at the edge of banana plantation 
east of bay, in leaf litter, leg. C. Schmidt 1.4. 1998; 2 immature males, 
I female (with marsupium): Caripe, surroundings of Cueva del 
Guacharo 10°10.37'N 63°33.14'W moist forest, under rotting loggs 
and leaf litter along brook, leg. C. Schmidt 7.4.1998; 1 male: 
Penfnsula de Paria. southern shore of eastern part, 10°34.87'N 
63°03.60'W, thermal well with thatched roof, uncovered near water 
edge: surrounded by meadows with cattle, leg. C. Schmidt 8.4.1998 

Ischioscia fasciifrons sp.nov. 

Figs l^t 

Material examined. Holotype, male (body length 10 mm): 
Curimagua. 1250 m. under wood in the garden of a hotel, leg. C. 
Schmidt 21.03.1998. 



Description 

Colour. Dorsum on a light umber ground colour patched with light 
spots and a darker medial band running down to the pleon, coxal 
plates with dark patches, on coxal plates VI and VII extensive dark 
areas. Cephalothorax with a dark band between the compound eyes 
on postfrons, profrons with a dark inverted Y covering lamina 
frontalis, extending ventrally. 

Cephalothorax, Vertex flattened with few tricorn-like setae, com- 
pound eyes consisiting of about 26 ommatidia arranged in four rows. 
Lamina frontalis and very faint linea supra-antennalis present, linea 
frontalis lacking (Fig. 1, Ctf). 

Pereon. Cephalothorax set back into pereonite 1 coxal plates 
prominent, with sulcus marginalis, no gland pores or noduli laterales 
visible in light microscope, tegument smooth, bearing several slen- 
der tricorn-like setae, highest density along lateral and distal margins 
(Fig. l,Cx3). 

Pleon. Set apart from pereon, neopleurae of pleonites 3 to 5 
prominent, pleotelson with concave lateral margin, as long as 
protopodites of uropods. 

Antennula. Composed of three articles of subequal length, distal 
article pointed with several aesthetascs apically and along medial 
margin, caudal side of proximal article extended (Fig. 1, Anl). 
Antenna. Broken in the specimen examined, peduncular articles 
bearing tricorn-like setae (Fig. 1, An2). 

Mandible. Pars intermedia with coniform setae, two penicils on 
left and one on right side, additional plumose seta proximally, molar 
penicil composed of about 7 branches (Fig. 2, Mdl/r). 
Maxillula. Medial endite with two stout penicils apically and 
small subapical tip, lateral endite with 4+6 teeth apically, 3 of inner 
set deeply, two slightly cleft, slender subapical tooth and acute 
hyaline lobe caudally (Fig. 2, Mxl). 

Maxilla. Both lobes covered with faint setation, medial trichiae 
stronger, lateral lobe slightly broader than medial, medial lobe 
apically cuspidate (Fig. 2, Mx2). 

Maxilliped. Basipodite with plentiful cover of tricorn-like setae, 
sulcus lateralis present, endite caudally setose, rostrally with knob- 
like penicil, palp with proximal article bearing short and long seta, 
medial article with short proximal setal tuft, distal tufts slightly 
stalked (Fig. 2, Mxp). 

Pereopods. Pereopods slender (Fig. 3, PE3-7; Fig. 4, PE1 ), carpus 
with antenna-grooming brush and ornamental sensory spine with 
hand-like apex (Fig. 3, Scl), dactylus with long inner claw and 
simple dactylar seta, apically slightly tapered (Fig. 3, Dae). Sexual 
differentiation. Pereopod 1 and 2 carpus enlarged with setal field 
rostrally, enlargement not too prominent, pereopod 3 with a very 
small setal field on medio-proximal border of caprus and medio- 
distal margin of merus. Pereopod 6 ischium with a small depression 
rostrally near medial margin with sensory spine, pereopod 7 ischium 
with a slightly twisted medial margin, thus forming a groove rostrally, 
slight distal lobe with few trichiae. 

Pleopods. Pleopod endopodites bilobate, exopodites with about 8 
to 9 sensory spines laterally, medially with trichiae and minute 
pectinate scales, pleopod 5 with caudal creel of three rows of 
prominent pectinate scales, protopodite 3 to 5 with medial protru- 
sion, less conspicuous on pleopod 4 and 5, rudimental epipodites on 
pereopod 1 to 4 (Fig. 4, PL1-5). Sexual differentiation. Male 
pleopod 1 exopodite triangular with lateral protrusion hardly set 
apart, incision merely a sinuosity, endopodite apically obtuse, with 
minute spines apically and lateral row of minute spines. Pleopod 2 
exopodite elongate with about 12 sensory spines laterally, medially 
with minute pectinate scales, endopodite longer than exopodite, 
apex looking like a hypodermic needle. 
Uropod. As in other species of the genus. 






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Fig. 1 Ischiosciafasciifrons sp.nov. Holotype, male, 10 mm. Anl antennula; An2 antenna; Ctf cephalothorax in frontal view; Cx3 coxal plate 3; Had 
habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 



140 



A. LEISTIKOW 




Mxl 



200nm 



Hy,. 2 hchioscia fasciifivns sp.nov. Holotype. male. 10 mm. Mdl/r left and right mandible; Mxl maxillula with detail of apex of lateral endite in rostral 
view; Mx2 maxilla in caudal view; Mxp maxilliped, with detail of endite in rostral view. 



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Fig. 3 Ischioscia fasciifrons sp.nov. Holotype, male, 10 mm. Dae dactylus 1 in rostral view; PE3-7 pereopods 3 to 7 in caudal view, ischium 7 in rostral 
view; Scl ornamental sensory spine of carpus 1, Spl distalmost sensory spine of propus 1. 



142 



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Ull- 4 1st Most in fasciifrons Sp.nov. Holotype. male. 10 mm. Gen genital papilla; PE1 pereopod 1 in rostral view; PL1-5 plcopods 1 to 5 in rostral view, 
with detail (it endopodite 1 in caudal view. 



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Genital papilla. Ventral shield slightly surpassing terminal spatula 
(Fig. 4, Gen). 

ETYMOLOGY . The species name refers to the conspiciously darker 
profrons, compared to the colouration of the cephalothorax. 

Ischioscia hirsuta sp.nov. 

Figs 5-8 

Material examined. Holotype, male (body length 11mm): An- 
des, Mesa cerrada between La Puerta and Timotes, 9°00.26'N 
70°44.20'W 1 800 m +/- 200 m, on the bank of a brook under stones 
and moist leaf litter, sugar cane, shrub with climbing plants, Polygon- 
aceae, Poaceae, leg. C. Schmidt 23.03.1998; Paratypes: 14males, 10 
females (with marsupium), 6 females, 2 immatures: same data as 
holotype; 6 males, 19 females (with marsupium), 15 females/ 
immatures: Andes, road from Timotes to Merida, 8°53.72'N 
70°47.99'W 3400 m (+/- 500 m), very steep northern slope, covered 
with Bryophyta, Pteridophyta, Poaceae, Ericaceae, between Bryo- 
phyta, leg. C. Schmidt 23.03. 1998 (MNHG); 3 males, 2 females (with 
marsupium), 1 females, 2 immatures: Andes, 8°43 . 1 2'N 70°46.04'W, 
on the bank of a brook near cattle meadow, srcub with Asteraceae, 
Rubus, one specimen submerged, leg. C. Schmidt 24.03.1998 

Description 

Colour. Ground colour chestnut with many light markings on 
cephalothorax and pereonites, coxal plates of same colour, dark 
brown band in the medial line of each pereonite, pleon bearing some 
light spots. 

Cephalothorax. Linea frontalis lacking, lamina frontalis and linea 
supra-antennalis prominent. Vertex smooth with plenty flagelliform 
tricorn-like setae, laterally protruding compound eyes composed of 
22 ommatidia in four rows (Fig. 5, Ctf). 

Pereon. Tegument smooth and shiny, bearing many flagelliform 
tricorn-like setae, coxal plates I to IV caudally rounded, coxal plates 
V to VII increasingly pointed, sulcus marginalis present, no gland 
pores nor noduli laterales discernible in light microscope at 400x 
magnification (Fig. 5, Cx4). 

Pleon. Set back from pereon despite neopleurae of pleonite 3 to 5 . 
Pleotelson rather pointed, lateral margins concave, some tricorn- 
like setae near the apex. 

Antennula. Similar to other species of genus (Fig. 5, Anl). 
Antenna. Flagellum three-articulate, with proximal article long- 
est, somewhat shorter than pedunclular article 5, apical organ longer 
than distal article, length ratio of peduncular articles similar to next 
species (Fig. 5, An2). 

Mandible. Pars intermedia bearing coniform setae and two penicils 
on left, one on right side, additional plumose seat proximally, molar 
penicil consisting of about 7 branches (Fig. 6, Mdl/r). 
Maxillula. Medial endite with two stout penicils apically and a 
short subapical tip, lateral endite with 4+6 teeth apically, 5 of inner 
set cleft, on caudal side, a hyaline lobe, stalk and two subapical teeth 
present (Fig. 6, Mxl). 

Maxilla. Lateral lobe slightly broader than medial, bearing faint 
trichiae and pectinate scales, lateral area of rostral side seta-free, 
medial one densely covered with trichiae, medial setal tuft present, 
apically cuspidate (Fig. 6, Mx2). 

Maxilliped. Basipodite with sulcus lateralis and many tricorn-like 
setae, palp with proximal setal tuft small, medial one stalked, as 
prominent as distal one, proximal article bearing a long and a very 
short seta, endite covered with trichiae, on rostral side with knob- 
like penicil (Fig. 6, Mxp). 

Pereopods. Slender with many sensory spines on medial margin 
(Fig.7, PE1-7), pereopod 1 carpus with antenna-grooming brush. 



propus with antagonistic device, dactylus with long inner claw, 
dactylar seta apically plumose (Fig. 8, Dae). Sexual differentiation. 
Pereopods 1 to 3 with subequally enlarged carpus and setal field 
rostrally, merus 1 to 3 with medial setal field, propus 2 with small 
setal field. Pereopod 7 ischium with prominent setal brush on 
proximal half medially, very long trichiae inserted on the more 
rostral surface, distally connected with a small cuticular clasp, 
bordered by small depressions, caudally a row of small trichiae, 
medio-distally with a lobe covered with short trichiae, laterally two 
sensory spine, third spine more caudodistally, basis 7 medio-distally 
with setal field around distal sensory spine. 

Pleopods. Pleopod endopodites slightly bilobate, exopodites with 
about 10 sensory spines laterally, pleopod 5 with creel of three rows 
of pectinate scales caudally, protopodites 3 to 5 with medial protru- 
sion, protopodites 4 and 5 lacking rudimental epipodite (Fig. 8, 
PL1-5). Sexual differentiation. Male pleopod 1 exopodite triangular 
with rounded medial edge, lateral point small, incision proxmally 
bordered by small protrusion, endopodite slender with lateral row of 
spines, apex with prominent tooth rostrally, faint trichiae caudally, 
medial protrusion subapically. Pleopod 2 exopodite as in other 
species, with minute pectinate scales medially, endopodite pointed. 
Uropod. As in other members of the genus. 
Genital papilla. Ventral shield slender, terminal spatule not sur- 
passing ventral shield (Fig. 8, Gen). 

Etymology. The species is named for its long tricorn-like setae 
on the pereonites, giving it a hairy appearance. 

Ischioscia colorata sp.nov. 
Figs 9- 12 

Material examined. Holotype, male (body length 14 mm): 
Aragua, Parque Nacional Henri Pittier, Road from Rancho Grande 
to the seaside, near Capilla Virgen del Carmen, mist forest, near 
brook and cascade in moist leaf litter and under stones, leg. C. 
Schmidt 14.03.1998. Paratypes, 10 females (with marsupium), 2 
females, 2 immatures: same data as holotype; 6 males, several 
immature males, 8 females, 4 immatures: Parque Nacional El Avila, 
10°33.01' N 66°53.88' W, high in the mountains at 1700 m (+/- 200 
m), within leaf litter near cliff, leg. C. Schmidt 10.03.1998. 

Description 

Colour. Pereonites chestnut with coxal plates slightly darker, many 
light spots of the muscle insertions and other light areas on coxal 
plates and neopleurae conspicuous, umber medial line, cephalothorax 
chestnut with light patches. 

Cephalothorax. Like other Venezuelan species of this genus, linea 
frontalis reduced, linea supra-antennal conspicuous, slightly bent 
above antennal sockets, compound eyes large, comprising about 26 
ommatidia, vertex bearing several long tricorn-like setae (Fig. 9, Ctf). 
Pereon. Tergites smooth and somewhat shiny, sparsely covered 
with tricorn-like setae, coxal plates fused without groove. Coxal 
plates 1-3 caudally rounded, 4-7 increasingly pointed. No noduli 
laterales visible in light microscope at 400x magnification (Fig. 9, 
Cx4). 

Pleon. Set back from the pereon, neopleurae of pleonite 3-5 well- 
developed. Pleotelson pointed with latero-distal margins concave. 
Tip of telson tapered, bearing tricorn-like setae of various length. 
Antennula. Three-articulate, first article strongest, with distal 
shield, distal article coniform, bearing at least 5 rows of aesthetascs 
(Fig.9, Anl). 

Antenna. Flagellum composed of three articles, proximal article 
of half the lentgh of flagellum, distal one shorter, bearing slightly 
longer apical organ. Peduncular articles 4 and 5 almost subequal. 



144 



A. LEISTIKOW 




Fig. 5 Ischioscia hirsute sp.nov. Holotype. male 1 1 mm. Aril antennula; An2 antenna with detail of apical organ; Ctfcephalothorax in frontal view; Cx4 
coxal plate 4; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 



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Fig. 6 Ischioscia hirsuta sp.nov. Holotype, male, 1 1 mm. Mdl/r left and right mandible; Mxl maxillula with detail of apex of lateral endite in rostral view; 
Mx2 maxilla in rostral view; Mxp maxilliped, with detail of endite in rostral view. 



146 



A. LEISTIKOW 




Spl Scl 



Fig. 7 l\i hioscia hirsuta Sp.nov. Holotypc. male, 1 1 mm. PE1 pcreopod I in rostral view: PES pcreopod 5 merus and ischium in caudal view; PE6/7 
pcrcopods IS and 7 in caudal view, with ischium 7 in rostral view; Scl ornamental sensory spine of carpus 1. Spl distalmost sensory spine ofpropus 



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147 




Fig. 8 lschioscia hirsuta sp.nov. Holotype, male, 1 1 mm. Dae dactylus 1 in rostral view; Gen genital papilla; PL1-5 pleopods 1 to 5 in rostral view, with 
detail of endopodite 1 in caudal view. 



I4S 



A. LEISTIKOW 




Fiu. <> hi hioscia colorata sp.nov. Holotype, male. 14 mm. Anl antennula; An2 antenna with detail of apical organ; Ctf'cephalothorax in frontal view; 
Cx4 coxal plate 4; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 



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Fig. 10 Ischioscia colorata sp.nov. Holotype, male, 14 mm. Mdl/r left and right mandible; Mx 1 maxillula with detail of apex of lateral endite in rostral 
view; Mx2 maxilla in rostral view; Mxp maxilliped, with detail of endite in rostral view. 



150 



A. LEISTIKOW 




Fig. 1 1 l\< hioscia colorata sp.nov. Holotype, male, 14 mm. Dae daetylus 1 in rostral view; PE4-7 pereopods 4 to 7 in caudal view, ischium 7 in rostral 
view;Scl ornamental sensory spine of carpus I : Si6 sensory spine of ischium 6; Spl distalmost sensory spine of propus 1. 



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Fig. 12 Ischioscia colorata sp.nov. Holotype, male, 14 mm. Gen genital papilla; PE1 pereopod lin rostral view; PL1-5 pleopods 1 to 5 in rostral view, 
with detail of endopodite 1 in caudal view. 



152 



A. LEISTIKOW 



each as long as flagellum, articles 2 and 3 half the length, proximal 
article short 'til covered with sensors spines and tricorn-like setae 
(Fig.9,An2). 

Mandible. Molar penicil with about 7 branches, pars intermedia 
with coniform setae, and two penieils on left, one on right side, 
additional plumose seta proximally (Fig. 10, Mdl/r). 
Maxillnla. Medial endite with two stout penieils and small subapi- 
cal tip. lateral endite with 4+6 teeth apically, 5 of inner set cleft, one 
of those double-cleft, two subapical teeth caudally. hyline lobe very 
slender (Fig. 10, Mxl). 

Maxilla. Lateral endite broader than medial, covered with pecti- 
nate scales, medial lobe with denser setation, apex with about 15 
cusps, setal tuft individualized (Fig. 10, Mx2). 
Maxilliped. basipodite with sulcus lateralis and many tricorn-like 
seate. palp with medial and proximal setal tuft stalked, proximal 
more indistinctively. proximal article with long and very short seta 
(Fig. 10, Mxp). 

Pereopods. Slender, as in other species of the genus (Fig. 11, 
PE4-7: Fig. 12. PE1), dactylus with long inner claw and apically 
plumose daetylar seta (Fig. 1 1, Dae), antenna-grooming devices on 
propus and carpus 1 (Fig. 12, PE1). Sexual differentiation. All 
pereopods sexually differentiated: pereopod 1 to 3 with enlarged 
carpus bearing setal fields rostrally, enlargement gradually de- 
creasing from 1 to 3, merus and propus with setal brushes, too. 
Pereopod 4 with scattered setae on medial margin of carpus and 
merus, pereopod 5 with setal field medio-proximally on merus, 
ischium with four stout sensory spines medially. Pereopod 6 merus 
with prominent hump medio-proximally, ischium with two stout 
sensory spines medially, pereopod 7 merus with long proximal 
sensory spine and four along medial margin, ischium as in other 
species in the variegata-group, i. e. a prominent medial brush, a 
distal lobe and a rostral depression. 

Pleopods. Pleopod endopodites slightly bilobate, exopodites with 
about 10 sensory spines laterally, medially distinctly covered with 
trichiae, exopodite 5 with creel of three rows of pectinate scales, 
distal ones parallel, proximal one diverging, protopodite 3 to 5 with 
slightly setose medial protrusion, laterally no rudimental epipodites, 
only few trichiae (Fig. 12, PL 1-5). Sexual differentiation. Male 
pleopod 1 exopodite obtuse triangular, lateral point recurved, inci- 
sion with transverse folding proximally on caudal side, endopodite 
straight, apex obtuse, rostrally with 5 to 6 teeth, caudally with tuft of 
hyaline trichiae, lateral row of spines present. Pleopod 2 exopodite 
elongate with sinuous lateral margin bearing 1 1 sensory spines, 
medially with trichiae, endopodite with truncate apex, bearing some 
faint granules caudally. 

Uropod. Exopodite and endopodite subequal in length. 
Genital papilla. Ventral shield slightly surpassing terminal spatula 
(Fig. 12, Gen). 

ETYMOLOGY. The specific name refers to the colourful dorsal 
tegument. 

Ischioscia pariae sp.nov. 

Figs 13-16 

MATERIAL EXAMINED. Holotype, male (body length 10 mm): 
Peninsula de Paria, El Refugio de la Cerbatana, primary forest 
with high trees covered with Brypohyta, Bromeliaceae, Lyco- 
podiaceae, vines, very humid, in leaf litter, leg. C. Schmidt 
04.04.1998. Paratypes: 4 males, 6 females, 7 immatures: same 
data as holotype: 7 males, 7 females (with marsupium), 8 females: 
Peninsula de Paria, El Rincon, 10°35.94' N 63°11.81' W, small 
brook, water only in upper half, humid forest, lower part in coacoa 



plantation, with some very high Bombacaceae, near small well, 
leg. C. Schmidt 09.04. 1998. 

Description 

Colour. Basic colour chestnut, prominent light patches on medial 
line of tergites and coxal plates, between them smaller irregular 
patches, caudal patches on median line dark umber, pleon chestnut 
with white patches in medial line, cephalothorax covered with small 
white spots, representing muscle insertions. 

Cephalothorax. Linea supra-antennalis prominent, linea frontalis 
lacking, with lamina frontalis, vertex flat, covered with some setae, 
compound eyes consisting of 25 ommatidia in 4 rows (Fig. 13, Ctf). 
Pereon. Tegument rather smooth and shiny, coxal plates 1 to 3 
rounded, coxal plates 4 to 7 caudally pointed, no noduli laterales nor 
gland pores discernible, sulcus marginalis present. 
Pleon. Retracted from pereon, pleonites 3 to 5 with small 
neopleurae, pleotelson with concave margins, densely covered with 
small cuticle-covered sensilla. 

Antennula. As in other species of genus, differing in position of 
aesthetascs on distal article, which are inserted at almost entire 
medial margin (Fig. 13, Anl). 

Antenna. Peduncle rather long with typical length ratio of other 
members of the genus, flagellum with proximal article longest, of 
half length of flagellum, apical organ longer than medial article (Fig. 
13, An2). 

Mandible. Molar penicil composed of about 7 branches, additional 
plumose seta long, pars intermedia with two penieils on left and one 
on right mandible, coniform setae on both sides (fig. 14, Mdl/r). 
Maxillula. Medial endite with two stout penieils and small sub- 
apical tip, lateral endite bearing 4+6 teeth apically, five of inner set 
cleft, on caudal side with hyaline lobe, stalk and two slender 
subapical teeth, laterally fringed with trichiae (Fig. 14, Mxl). 
Maxilla. Lateral lobe slightly broader than medial one, covered 
with pectinate scales and faint trichiae, medial lobe densely covered 
with trichiae, apically with about 15 cusps (Fig. 14, Mx2). 
Maxilliped. Basipodite with sulcus lateralis, palp with one seta on 
proximal article, medial article with two setal tufts, distal one stalked, 
distal article with prominent setal tuft, endite caudally setose with 
two teeth, rostrally with prominent penicil (Fig. 14, Mxp). 
Pereopods. Pereopod 1 carpus and propus with antenna-grooming 
brushes (Fig. 15, PE2-7; Fig. 16, PE1), dactylus with a hyaline 
cuticular scale laterally of main claw, inner claw long, daetylar seta 
with an enlarged apex, subapically some plumules (Fig. 15, Dae). 
Sexual differentiation. Male pereopods 1 to 3 with enlarged carpus 
and prominent setal fields rostrally, carpus 2 and 3 only slightly 
enlarged, merus 1 to 3 with medial setal field, setal field present on 
propus 2, too. Pereopod 7 ischium with distal lobe, sparsely covered 
with trichiae, setal brush on medio-proximal half slightly directed 
rostrodistally, more laterally accompanied by a prominent groove 
extending to the lateral margin, two sensory spines on lateral 
margin. 

Pleopods. Endopodites more rectangular and exopodites more 
triangular as in preceding species, especially in pleopod 5, laterally 
bearing 6 to 8 sensory spines, protopodite 3 to 5 lacking rudimental 
epipodites (Fig. 16, PL1-5). Sexual differentiation. Pleopod exopodite 
triangular with rounded medial edge and slighly sinuous medial 
margin, almost continued by lateral protrusion, incision proximally 
bordered by a step-like additional tip, endopodite straight, apically 
pointed with small lateral protrusion subapically, caudolateral row of 
minute spines, single prominent spine apically. Pleopod 2 with only 
slightly sinuous lateral margin bearing 8 sensory spines, endopodite 
with apex shaped like a hypodermal needle. 
Uropod. Exopodites rather long (Fig. 13, UR). 



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153 




lOOum 



Fig. 13 Ischioscia pariae sp.nov. Holotype, 13 mm. Anl antennula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; Had 
habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson; UR uropod in rostral view. 



154 



A. LEISTIKOW 




Fig. 14 Ischioscia pariae Sp.nov. Holotype, 13 mm. Mdl/r left and right mandible; Mxl maxillula in caudal view, with detail of apex of lateral endite in 
rostral view; Mx2 maxilla in rostral view: Mxp maxilliped. with detail of endite in rostral view. 



NEW SPECIES OF ISCHIOSCIA FROM VENEZUELA 



155 




Fig. 15 Ischioscia pariae sp.nov. Holotype, 13 mm. Dae dactylus 1 in rostral view; PE2-7 pereopods 2, 6, 7 in caudal view, ischium 7 in rostral view; Scl 
ornamental sensory spine and proximal sensory spine of carpus 1; Spl distalmost sensory spine of propus 1. 



156 



A. LEISTIKOW 




Fig. 16 hchioscia pariae sp.nov. Holotype, 13 mm. Gen genital papilla; PEI pcrcopod lin rostral view; PL1-5 pleopods 1 to 5 in rostral view, with 
detail of endopodite I in caudal view. 



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157 



Genital papilla. Ventral shield distinctly surpassing terminal 
spatula (Fig. 16, Gen). 

Ecological note. This species is capable of jumping about 5 cm. 

ETYMOLOGY. The species name refers to the distribution on 
Peninsula de Paria. 



Ischioscia guamae sp.nov. 

Figs 17-20 

Material examined. Holotype, male (body length 1 3 mm): Rio 
La Guama, 9°46.48' N 68°24.6' W, river valley between dry moun- 
tains, in leaf litter, leg. C. Schmidt 25.03.1998. Paratypes 1 males, 3 
females llimmatures: same data as holotype. 

Description 

Colour. Ground colour chestnut with several light markings on 
cephalothorax and pereonites, coxal plates with light margins, in 
coxal plates V to VII, two light patches instead of contiguous band, 
dark chestnut spot in the mid line of pereonite I to IV caudally, pleon 
bearing several pale spots. 

Cephalothorax. Linea frontalis lacking, lamina frontalis and linea 
supra-antennalis prominent. Vertex smooth and free of setation, 
laterally protruding compound eyes composed of 22 ommatidia in 
four rows (Fig. 17, Ctf). 

Pereon. Tegument smooth and shiny, bearing scattered setae, coxal 
plates 1 to 4 caudally rounded, coxal plates 5 to 7 increasingly pointed, 
sulcus marginalis present, no gland pores nor noduli laterales discern- 
ible in light microscope at 400x magnification (Fig. 1 7, Cx3). 
Pleon. Set back from pereon despite neopleurae of pleonite 3 to 5. 
Pleotelson rather pointed, lateral margins concave, some tricorn- 
like setae apically. 

Antennula. Similar to other species of genus (Fig. 17, Anl). 
Antenna. Rather short, flagellum three-articulate, with proximal 
article longest, somewhat shorter than pedunclular article 5, tricorn- 
like setae on all articles, apical organ half as long as flagellum (Fig. 
17, An2). 

Mandible. Pars intermedia with two penicils on left and one on 
right side, several coniform setae, molar penicil composed of about 
7 branches, additional plumose seta prominent (Fig. 18, Mdl/r). 
Maxillula. Medial endite with two stout penicils and very small 
subapical tip, lateral endite apically with 4+6 teeth, 4 of inner set 
cleft, 2 very slender, caudally with hyaline lobe and slender subapi- 
cal tooth (Fig. 18, Mxl). 

Maxilla. Lateral lobe boader than medial lobe, covered with faint 
trichiae and pectinate scales, medial lobe densely setose, apically 
with aboput 15 cusps (Fig. 18, Mx2). 

Maxilliped. Palp with three setal tufts medially, distal two very 
prominent, proximal article bearing long and short seta, endite with 
knob-like penicil on rostral side, caudally setose with two teeth, 
basipodite with sulcus lateralis (Fig. 18, Mxp). 
Pereopods. Slender(Fig. 19,PEl-7),withdactylusbearingplumose 
dactylar seta and long inner claw (Fig. 19, Dae), pereopod 1 carpus 
with antenna-grooming brush and ornamental sensory spine with 
hand-like apex (Fig. 1 9, Sc 1 ), propus 2 with antenna-grooming comb. 
Sexual differentiation. Pereopod 1 to 3 and 7 differentiated, propus 1 
and 2, carpus and merus 1 to 3 with setal brush, carpus medially 
enlarged, conspicuously in pereopods 1 to 2, slightly in pereopod 3. 
Pereopod 7 ischium with setose medio-distal lobe, setal brush 
subproximally to halflength, no rostral depression, merus 7 of charac- 
teristic shape, 2 medio-caudal sensory spines on proximal half. 
Pleopods. Similar to other species of the genus, exopodites later- 
ally with sensory spines, medially with minute pectinate scales, no 



rudimental epipodites on protopodites 3 to 5 (Fig. 20, PL 1-5). 
Sexual differentiation. Male pleopod 1 exopodite triangular with 
lateral incision bordered by lateral point and proximal protrusion, 
endopodite with wrinkeled apex and subapical lateral protrusion, 
caudal row of spines reduced, some spines on apex, two of them very 
prominent, surpassing apex. Pleopod 2 endopodite slighly sinuous 
laterally, bearing about 7 sensory spines, endopodite slighly sur- 
passing exopodite, apex pointed. 
Uropod. As in other species of the genus. 

Genital papilla. Ventral shield slighly surpassing terminal spatula 
(Fig. 20, Gen). 

Etymology. The species name is derived from Rio La Guama, 
the type locality. 

Ischioscia trifasciata sp.nov. 

Figs 21-24 

Material examined. Holotype, male (body length 13mm), 
Rancho Grande, 10°21'N 67°41'W, in ground traps 30 cm diameter, 
leg. O. Hernandez 1995. Paratypes: several males and females: same 
data as holotype. 

Description 

Colour. Dorsal tegument rich chestnut with many yellowish 
patches, medial line and coxal plates dark umber brown, forming 
three bands, margin of coxal plates lighter, pleon chestnut with pairs 
of light spots on each pleonite, cephalothorax chestnut, vertex 
heavily spotted yellowish. 

Cephalothorax. As in other species of Ischioscia with large, later- 
ally protruding compound eyes composed of about 24 ommatidia, 
vertex somewhat flattened, linea frontalis lacking, linea supra- 
antennalis conspicuous, only slightly sinuous, between antennal 
sockets with prominent lamina frontalis (Fig. 21, Ctf). 
Pereon. Coxal plates of pereonite 1-3 with rounded margins, 
fourth to seventh coxal plate increasingly caudally pointed, no 
noduli laterales. Tegument smooth with only few tricorn-like setae 
(Fig. 21, Cx4). 

Pleon. Retracted from pereon, pleonites 3-5 with small neopleurae. 
Pleotelson with concave latero-distal margins, as long as protopodites 
of uropods, tricorn-like setae gathered at apex. 
Antennula. Apex of three-articulate antennula coniform, termi- 
nated by tuft of three aesthetascs, row of at least four aestethascs on 
medial border. Some hairlike setae on median article (Fig. 21, Anl). 
Antenna. Peduncular articles with various sensory and tricorn-like 
setae as dense as on flagellum. Fifth article the longest, fourth 3/4 the 
length of fifth, as long as third and second together, those subequal. 
Flagellar articles subsequently shorter, together longer than fourth 
peduncular article, apical organ of same length as distal flagellar 
article (Fig. 21, An2). 

Mandible. Pars intermedia with two stout penicils on left, one on 
right mandible, proximally additional penicil, molar penicil com- 
posed of 6 to 7 branches (Fig. 22, Mdl/r). 

Maxillula. Medial endite with 2 stout penicils apically, small 
apical tip, lateral endite laterally fringed with trichiae, apically 
bearing 4+6 teeth, 5 of inner set cleft, caudally with hyaline lobe, 
stalk and two small teeth with fringed apex (Fig. 22, Mxl). 
Maxilla. Both lobes subequal in breadth, densely covered with faint 
trichiae, medial lobe apically with about 12 cusps (Fig. 22, Mx2). 
Maxilliped. Basipodite with sulcus lateralis, endite with tooth and 
dense setation caudally, rostrally with conspicuous knob-like penicil, 
palp with proximal article bearing two unequal setae, distal articles 
fused without a groove, three setal tufts of 7 to about 25 setae, 
proximal and medial stalked (Fig. 22, Mxp). 



158 



A. LEISTIKOW 




H«. 17 Ischioscia guamae Sp.nov. Holotype, male 13 mm. An I antcnnula; An2 antenna with detail of apical organ; Ctf cephalothorax in frontal view; 
Cx3 COXal plate 3; Had habitus in dorsal view; Hal habitus in lateral view; Tel pleotelson. 



NEW SPECIES OF ISCHIOSCIA FROM VENEZUELA 



159 




Fig. 18 Ischioscia guamae sp.nov. Holotype, male 13 mm. Mdl/r left and right mandible; Mxl maxillula with detail of apex of lateral endite in rostral 
view; Mx2 maxilla in rostral view; Mxp maxilliped, with detail of endite in rostral view. 



160 



A. LEISTIKOW 




Spl Scl 



Fie. 19 hchioscia guamae sp.nov. Holotype. male 13 mm. Dae clactylus 1 in rostral view; PE1 pereopod lin rostral view; PE3-7 pereopods 3, 6, 
7 in caudal view, ischium 7 in rostral view; Scl ornamental sensory spine of carpus 1; Spl distalmost sensory spine of propus 1. 



NEW SPECIES OF ISCHIOSCIA FROM VENEZUELA 



161 




PL2 



Fig. 20 Ischioscia guamae sp.nov. Holotype, male 13 mm. Gen genital papilla; PL 1-5 pleopods 1 to 5, rostral view, with detail of endopodite 1 in 
rostral and caudal view. 



162 



A. LEISTIKOW 




Fig. 21 Ischioscia trifasciata sp.nov. Holotype. male 13 mm. Anl antennula; An2 antenna with detail of apical organ: Ctf cephalothorax in frontal view; 
Cx4 eoxal plate 4; Had habitus in dorsal view; Hal habitus in lateral view: Tel pleotelson. 



NEW SPECIES OF ISCHIOSCIA FROM VENEZUELA 



163 




200nm 



Fig. 22 Ischioscia trifasciata sp.nov. Holotype, male 1 3 mm. Mdl/r left and right mandible; Mx 1 maxillula with detail of apex of lateral endite in rostral 
view; Mx2 maxilla in rostral view; Mxp maxilliped, with detail of endite in rostral view. 



164 



A. LEISTIKOW 




H>>. 23 Ischioscia trifasciata sp.nov. Holotype, male 1 3 mm. Dae dactylus 1 in rostral view; PE1 pereopod lin rostral view; PE5-7 pereopods 5 to 7 in 
caudal view, ischium 7 in rostral view; Scl ornamental sensory spine of carpus I ; Si6 sensory spine of ischium 6; Spl distalmost sensory spine of propus 1. 



NEW SPECIES OF ISCHIOSCIA FROM VENEZUELA 



165 




PL2 



PL5 



Fig. 24 Ischioscia trifasciata sp.nov. Holotype, male 1 3 mm. Gen genital papilla; PL 1-5 pleopods 1 to 5, rostral view, with detail of endopodite 1 
rostral and caudal view. 



166 



A. LEISTIKOW 



Pereopods, Slender and long (Fig. 23. PE1-7), dactylus with 
simple dactj lar seta, apicallj « ith some rudiments of plumes, long 
inner claw (Fig. 23, Dae), pereopod 1 carpus with antenna-grooming 

brush and ornamental sensor) spine with hand-like apex (Fig. 23, 
Scl), distal sensor\ spine of basis long. Sexual differentiation. Male 
pereopods I to 3 with seta! brushes on propus to merus, carpus 
enlargement subequal in all 3 pereopods, merus of pereopod 5 and 6 
with set o{ sensory spines of different length, proximal spine of 
merus 6 on a slight hump, direeted distally, pereopod 7 merus 
mediall} w ith proximal sensory spine, two long spines on halflength, 
two shorter ones more proximally. ischium with dense setal brush, 
medio-distal lobe with dense cover of small cutieular trichiae, 
rostrally with a slight depression. 

Pleopods. Pleopod endopodites slightly bilobate, exopodites with 
up to 10 sensory spines laterally and minute peetinate scales medi- 
ally, protopodites 3 to 5 without lateral rudiments of epipodites, no 
respiratory areas discernible (Fig. 24. PL 1-5). Sexual differentia- 
tion. Male pleopod 1 exopodite triangular with long lateral point, 
lateral margin straight, small wrinkle proximally of point, endopoite 
slender with short subapical protrusion laterally, apex drawn out 
with 5 spines rostrally, caudally with dense row of spines. Pleopod 
2 exopodite with sinuous lateral margin, endopodite with slightly 
pointed apex. 

Uropod. As in generic diagnosis. 

Genital papilla. Ventral shield only slightly surpassing terminal 
spatula (Fig. 24, Gen). 



DISCUSSION 

Most of the species of Ischioscia Verhoeff, 1928 found in Venezuela 
belong into the vicinity of the well-known /. variegata (Dollfus, 
1893). For a detailed description of this species cf. Leistikow 
(1997). /. fasciifrons sp.nov. is distantly related to the others; it 
differs in the lack of a setal brush on the male pereopod 7 ischium 
which bears a transverse furrow medio-rostrally instead of this 
brush, the distal lobe is rather inconspicuous. Another autapomorphy 
of this species is the simple dactylar seta instead of an apically 
plumose one. The apex of the male pleopod 1 endopodite is obtuse 
with a set of very small spines terminally, quite distinct from the 
medio-caudal row of spines, the exopodite is only slightly incised, 
with a short protrusion. The structure of the male pleopod 1 is similar 
to /. colorata sp.nov. from the coastal region of Caracas, this species 
differs in the presence of prominent setal fields in the carpus and 
merus of pereopod 4, merus of pereopod 5 and a setal brush on the 
ischium of pereopod 7. Thus. /. colorata is a member of the 
variegata-group of species as defined by Leistikow (1997). This 
species is further characterized by the shape of the apex of male 
pleopod 2 endopodite, which is an autapomorphy for the species. 
The maxillula bears a single subapical tooth like /. hirsuta sp.nov, 
and /. fasciifrons sp.nov., this is in contrast to the two teeth in /. 
patriae sp.nov.. /. gtiamae sp.nov., and /. trifasciata sp.nov. 

In /. trifasciata, both teeth are apically serrate while in the others 
the teeth are acute. Besides, the simplified dactylar seta, the shape of 
the male pleopod 1 and the setation of male merus 7 are autapo- 
morphies of/, trifasciata. 

I. hirsuta is characterized by the following derived character 
states: a long-haired setal brush on male ischium 7, a hand-like 
subdistal sensory spine of propus 1, and the long tricorn-like setae 
on the pereonites. 

For /. guamae, the most distinct character is the setation of the 
male merus 7. with the four sensory spines all in the proximal half of 



1+2+1+0 + 3 




Fig. 25 Schematic drawing of the male merus 7, indicating the position 
of sensory spines along medial margin in the ground pattern of the 
variegata-group within the genus Ischioscia Verhoeff, 1928. 

the article, one of the smaller stands alongside the long proximal 
sensory spine. Another autapomorphic character is the caudolateral 
position of the remainder of the row of spines on the male pleopod 
1 exopodite. 

The autapomorphies for /. pariae are the peculiar shape of the 
dactylar seta with club-like apex instead of the feathery one, and the 
reduction of the number of sensory spines on the male merus 7 to 
three along the medial border. 

The setation of the male merus 7 allows easy recognition of the 
Venezuelan species of Ischioscia. In the ground pattern of the male 
merus 7, the arrangement of sensory spines along the medial margin 
from proximally to distally is a long proximal one, two smaller ones 
distally of the former, another sensory spine even more distally and 
then a gap which is terminated distally by the medio-distal set of 
about three long sensory spines, described by the formula 
[1+2+1+0+3] which can be found in e.g. /. martinae Leistikow, 
1997, /. plurimaculata Leistikow, 1999 or /. bolivari Vandel, 1968 
(fig. 25). This formula is varying in the different species of Ven- 
ezuela as follows: 



/. hirsuta 
I. colorata 
I. pariae 
I. guamae 
I. trifasciata 
I. variegata 



[l+(l+2)+(l+2)+l+3] 
[1+2+1 + 1+3], 
[(l + D+0+1+0+3] 
[(1 + 0+1 + 1+0+3] 
[1 + 1+3+0+3] 
[0+( 1 + 0+0+0+3]. 



I. fasciifrons does not fit this sheme as it differs in other characters 
as stated above. The position of the long spine alongside a single 
short one may be a synapomorphy of/, pariae and /. variegata, this 
assumption is supported by the similar shape of the genital papilla, 
which in both species has a long pointed ventral shield - much 
longer than the termial spatule with the laterally placed orifices. 

From an ecological point of view, the genus Ischioscia is adapted 
to many different types of habitat. The autochthonous habitat is the 
moist rain forest but the members of this genus have adapted to 
secondary growth, cattle pastures and other disturbed environments. 
An interesting behavioural difference has been observed in /. pariae 
and /. variegata both of which are capable of jumping. The former 
was observed to jump about 5 cm; /. variegata can jump even wider, 
up to 20 cm and hence can easily escape predators like spiders and 



NEW SPECIES OF ISCHIOSCIA FROM VENEZUELA 



167 




Fig. 26 Distribution of the species of Ischioscia Verhoeff, 1928 in Venezuela. □, I.guamae; ■£, /. pariae; ♦ . /. colorata; #, /. trifasciata; #. /. hirsuta; 
+, /. fasciifrons: O, /. variegata. 



even the astonished human collector. The strong pereopods 6 and 7 
with the powerful musculature in the basis are essential for this type 
of locomotion. The jumping capability may provide another 
synapomorphy for the two species. Another strategy to escape 
predators was observed in /. hirsuta. A specimen which was observed 
at the bank of a small brook fled under water and stayed submerged 
for a considerable period. 

These new records widen our knowledge on the genus particu- 
larly with respect to their biogeography. The genus is well-distributed 
in Costa Rica (Leistikow 1999), most records from Venezuela refer 
to the most abundant species /. variegata, which is distributed in the 
areas adjacent to the Caribbean Sea and also in the interior of 
Venezuela along the western Corilleras (Fig. 26). The other species 
seem to have more restricted ranges: /. pariae is confined to the 
Peninsula de Paria; /. colorata was found in the vicinity of Districto 
Federal. These patchy distributions may be collection artifacts. But 
they may reflect a characteristic biogeographical pattern found in 
many Neotropical taxa. Cracraft (1985) argued for the generality of 
areas of endemism in South America. It is possible to compare 
distributional data obtained from such diverse taxa as plants (Prance 
1982), butterflies (Whitmore & Prance 1986) or birds (Haffer 1974, 
Cracraft 1985). They all support distinct regions of high endemism 
called Pleistocene refuges by Haffer (1974). The distribution of 
species of Ischioscia in Venezuela also corresponds to those regions. 



/. pariae is found in the Parian centre, /. colorata in the Venezuelan 
Montane centre and I. hirsuta in the Meridan Montane centre. The 
widespread /. variegata might be an expansive species best adapted 
to human habitats, although some of the records may be erroneous 
(Arcangeli 1930, Richardson 1914). /. variegata may occur in close 
spatial vicinity of a second species of Ischioscia, but they were not 
encountered in exactly the same locality. Thus, effective isolation 
mechanisms have to be postulated to separate the species. 



Acknowledgments. The author is grateful to Dr. C. Schmidt for permis- 
sion to analyse the philosciid samples. Moreover, he is indebted to Dr. H. 
Schmalfuss, Staatliches Museum fur Naturkunde, for the loan of material of 
Ischioscia from the museum's collection. 



REFERENCES 



Arcangeli, A. 1930. Contribute alia conoscenza del 'Microgenton' di Costa Rica. 

Bollettino del Laboratorio di Zoolgia Agraria delta Faccolta Agraria 25: 1-29. 
Cracraft, J. 1 985. Historical biogeography and patterns of diversification in Neotropical 

birds. Ornithological Monographs 36: 49-84. 
Haffer, J. 1974. Avian speciation in tropical South America. Publications of the Nuttalt 

Ornithological Club 14: 1-196. 
Leistikow, A. 1 997. Terrestrial Isopods from Costa Rica and redescription of Ischioscia 

variegata (Dollfus, 1893). Canadian Journal of Zoology IS: 1415-64. 



168 



A. LEISTIKOW 



1999. New species of the genera Ischioscia and Andenoniscus, terrestrial isopods 

from Panama (Crustacea, Isopoda). Studies on neotropical Fauna and Environment 

34: 156 1 5 
21XX). The terrestrial isopod genus Ischioscia in Costa Rica: new species and 

records, andananalysisof its ph> logenv (Crustacea: Isopoda: Oniscidca). Af(rre , //wi#<'M 

aus dem Museum tur Xalitrkiunle in Berlin, Zoologische Reihe 76 ( I ): 1—49. 
I.emns de Castro, A. 1955. Ischioscia amazonica, uma nova especie de isopode 

tenestre do Estado do Amazonas (Isopoda: OmsciA&t). Revistabrasileirade Biologic 

15: 1 S 
Prance, G.T. 1982. Fores) refuges: Evidence from woody angiosperms. pp. 137-150 

In: G.T. Prance (Ed) Biological diversification in the Tropics. New York. 
Richardson. H. 1914 Tenestrial isopods of Colombia. Memoires de la Societe de 

\, uch&tel Sciences Nalurcllcs 14: 29-32. 
Schmalfuss, H. 1980. A revision of the neotropical genus Ischioscia Verhoet'f. with 



description of four new species (Isopoda. Philosciidae). Studies on neotropical 

Fauna and Environment 15: 125-139. 
Vandel, A. 1968. Isopodes terrcstres. In: N. and J. Leleup (Eds) Mission zoologique 

beige aux lies Galapagos el Ecuador, 2 (84): 35-168. 
1972. Les isopodes terrestres de la Colombie. Studies on neotropical Fauna and 

Environment 7: 147-172. 
van Name, W.G. 1926. Forest isopods from Barro Colorado Island, Panama Canal 

Zone. American Museum Novitates 106: 1-15. 
1 936. American terrestrial and fresh water Isopoda. Bulletin of the American 

Museum of Natural History 71: 1-520. 
Verhoeff, K.W. 1928. Uber einige Isopoden der zoologischen Staatssammlung in 

Munchen. Zoologischer Anzeiger 75: 25-361. 
Whitmore, T.C. & Prance, G.T. 1986. Biogeography and Quaternary history in 

tropical America. 212p. Oxford Univerity Press, Oxford. 



Bull. nat. Hist. Mus. Lond. (Zool.) 67(2): 169-182 



X* (3<vo<b£^ ,\J 



Issued 29 November 2001 



A review of the genus Erenna Bedot, 1904 
(Siphonophora, Physonectae) 



RR. PUGH 

Southampton Oceanography Centre, Empress Dock, Southampton S014 3ZH 



SYNOPSIS. The status of the poorly known physonect genus Erenna is reviewed in the light of the collection, by submersibles. 
of specimens in excellent condition. The few previous descriptions had been based on only the tentacles, or on other parts in poor 
condition. Two species have been described, E. richardi Bedot, 1904 and E. bedoti Lens & van Riemsdijk, 1908, and there has 
been some debate as to whether they are conspecific or not. It is concluded here that they are conspecific. Two further Erenna 
species, E. laciniata sp.nov. and E. cornuta sp.nov., are described, together with a third, closely related species, Parerenna 
emilyae sp.nov. The distinctiveness of their tentilla, with uncoiled hypertophied cnidobands, and the nectophores with a basic 
ridge pattern and a muscle-free zone at the apex of the nectosac, is considered sufficient to warrant the transference of these 
species into a new physonect family, the Erennidae. 



INTRODUCTION 



Very little is known about the physonect genus Erenna, and speci- 
mens have rarely been identified. In fact, the first species, E. 
richardi Bedot, 1904, was originally described from only six tenta- 
cles found attached to the rope of a fish trap brought up from a depth 
of 5310 m somewhere between Portugal and the Azores. However, 
these tentacles bore highly distinctive tentilla of a type not previ- 
ously known for any siphonophore, with uncoiled, hypertrophied 
cnidobands (stinging bands), and with the gastrodermal walls of the 
gastrovascular cavity packed with dark granules. Bedot (1904) 
made a detailed histological study of these tentilla and suggested 
that they probably belonged to a physonect siphonophore. 

A second species, Erenna bedoti Lens & van Riemsdijk, 1908, 
was described from a denuded specimen collected during the Siboga 
Expedition. From the fragmented material, with many appendages 
containing black pigmentation, Lens and van Riemsdijk (1908) 
deduced that the species was a physonect siphonophore, and they 
assigned it to the family Forskaliidae. Although they could compare 
only the tentacles and tentilla with Bedot's (1904) material, they 
reasoned that their material could be referred to a different species 
because the free end of the cnidoband, on the more developed 
tentilla in their material, lay proximally, whereas it was distal in E. 
richardi. However, most subsequent authors have considered E. 
bedoti to be conspecific with E. richardi. 

Bigelow (1911, p. 271) mentioned another specimen of Erenna 
richardi, in poor condition: 'In fact the condition is so bad that it is 
impossible to state whether or not it is specifically identical with the 
'Siboga' example. Nor, for that matter, is it clear whether the latter 
is distinct from Bedot's E. richardi.' . However, he, like Lens and 
van Riemsdijk (1908), considered that its closest relatives were the 
forskaliids. Moser (1925) described a further specimen collected in 
the Bay of Biscay, which consisted of only a poorly preserved 
siphosome. She noted that the gastrozooids appeared not to have a 
pedicle (stalk), and that their gastrodermal lining contained black 
granules. On the basis of the structure of the gastrozooids and 
tentacles she considered that it could not be a forskaliid siphonophore 
and she likened it to a bathyphysid (Order Cystonectae), but noted 
that the latter lacked nectophores. 

Tentacles, with characteristic tentilla were collected, in the north- 
east Atlantic on two further occasions (Leloup, 1936), but then no 



further specimens of Erenna species were recorded until Totton 
(1965) found three more. These were all in poor condition, but 
sufficient for the mature nectophores and bracts to be described and 
illustrated for the first time. The apex of the nectosac of the large, 
flattened nectophores was found to be muscle-free; and the radial 
canals could have more or less well-developed 'horn' canals ascend- 
ing into the mesogloea. The bracts were said to have two pairs of 
lateral processes. 

Margulis (1969) added to the description of the nectophores of 
Erenna richardi, noting that there were two small processes on the 
ventral side of the thrust block (the central region that abuts the 
stem) and that 'horn' canals were not always present on the lateral 
radial canals. In a later paper (Margulis, 1977) she described this 
specimen in more detail and concluded, from the shape of the mature 
tentilla, that it could be referred to Erenna richardi. However, she 
then briefly discussed possible differences from Lens and van 
Riemsdijk's (1908) specimen and concluded that E. bedoti was also 
a valid species. Margulis (1990) used these differences to describe a 
further specimen of E. bedoti collected in the southern Pacific. 
Recently, other specimens off. richardi have been briefly described 
and/or recorded (Pugh, 1975; Musayeva, 1976; Alvarino, 1980; 
Leloup, 1980; Daniel, 1985). 

In recent years, specimens of Erenna species have been collected 
by the submersibles Johnson-Sea-Link (JSL) I and II, and these are 
here used to give a more detailed description off. richardi, together 
with descriptions of two other, previously undescribed, species that 
can be referred to the same genus. In addition, another JSL speci- 
men, that is closely related to the genus Erenna, will be described. 
The taxonomic status of these species is discussed below and it is 
concluded that they should be separated off into a new family. The 
validity of E. bedoti also is discussed. 



Family ERENNIDAE fam. nov. 

Diagnosis. Physonect siphonophores best characterised by their 
uncoiled tentilla bearing a hypertrophied cnidoband with nematocy sts 
of three types: large anisorhizas and two types of smaller ones (? 
haplonemes) . Terminal process devoid of nematocy sts . Nectophores 
with basic ridge pattern of apico-, infra- and vertical laterals; with 
apical muscle-free zone on nectosac; radial canals straight or slightly 
curved. Ostium, without mouth plate, opening basally. 



© The Natural History Museum, 2001 



170 



P.R.PUGH 



Pneumatophore \\ ithoul apical pore. Gastrozooids without pedicle. 
Dioecious 

R] M VRKS. As noted in the Introduction, both Lens and van 
Riemsdijk ( 1908) and Bigelow (1911) considered the genus Erenna 
mighl be related to the physonect family Forskaliidae. whereas 
Moser 1 1925) associated it with bathyphysid cystonects. However, 
lotion 1 1965) placed Erenna in the physonect family Agalmatidae. 
As Pugh (1998) discussed, the Agalmatidae is probably a composite 
family containing all those species that do not have the distinctive 
characters of other physonect families. There is a core of similar 
genera. Agalma, Halistemma and Lycknagalma, which have 
involucrate tentilla. with lightly coiled cnidobands. Nanomiu is 
somewhat similar. However, the other genera are often difficult to 
relate to each other in any basic way. 

Pugh (1999) discussed this further, with regard to the genus 
Bargmannia, which Totton (1965) had placed in the family 
Pyrostephidae. Among the key characters used in establishing the 
taxonomic position of that genus were the ridge pattern on the 
nectophores; the presence of a muscle-free zone on the nectosac; and 
the structure of the tentillum. With regard to the ridge pattern, which 
consisted of pairs of apico-, infra and vertical (meso-) laterals, he 
noted that a similar arrangement was found on the nectophores of 
Pyrostephos vanhoeffeni Moser, 1925 and Erenna richardi; with an 
even simpler arrangement, omitting the vertical laterals, being 
found in Marrus species. Similarly, all these species had a muscle- 
free zone on the nectosac. In addition, Bargmannia species, P. 
vanhoeffeni and M. antarcticus were known to be dioecious, whereas 
most other physonects are known to be monoecious. In the present 
study it will be shown that two Erenna species are dioecious, with 
monovan gonophores; while gonophores were not found with the 
other two species. These characters separate these four genera from 
all other agalmatids. 

In Bargmannia spp., Pyrostephos vanhoeffeni and Marrus spp. 
the tentilla have simple, straight, or loosely coiled, cnidobands; with 
long contractile terminal filaments bearing nematocysts. However, 
Erenna richardi has a straight, hypertrophied cnidoband; and a rigid 
terminal process devoid of nematocysts. There are also differences 
in the types of nematocysts present on the tentillum. For many 
agalmatids four types are present: homotrichous anisorhizas 
(haplonemes), and either mastigophores or stenoteles on the 
cnidoband: desmonemes and acrophores in the terminal filament. 
The tentillum of Marrus species appears to conform with this 
pattern, with microbasic mastigophores included in the cnidoband. 
In Bargmannia species and P. vanhoeffeni large nematocysts, prob- 
ably stenoteles, were present only on the proximal part of the 
cnidoband: with two types of smaller nematocysts present through- 
out the remainder of the cnidoband and terminal filament. Pugh 
(1999) was uncertain whether the latter were acrophores or 
desmonemes, but haplonemes were thought to be absent. As is 
shown below, the cnidobands of Erenna species contain 
mastigophores and two types of haplonemes; but, as noted above, 
there are no nematocysts on the terminal process. 

Pugh (1999) concluded that there were sufficient similarities 
between Bargmannia spp. and Pyrostephos vanhoeffeni to retain the 
former in the family Pyrostephidae. However, despite the similari- 
ties of the nectophoral ridge pattern and the muscle-free zone on the 
nectosac. there are certain marked differences between these species 
and those of the genus Erenna. This particularly applies to the 
structure of the tentillum and its nematocysts; but also to the general 
structure of the nectophore. Pyrostephid nectophores have a large 
triangular thrust block and the axial wings are either reduced or 
absent; with the lateral radial canals on the nectosac arising separately 



from the dorsal canal. In Erenna spp. the thrust block is much 
smaller, while the apical wings are well-developed, and all four 
radial canals arise from the pedicular canal either together, or very 
nearly so. In addition, palpons with palpacles (reduced tentacles) are 
present in Erenna species, while they are totally absent in Bargmannia 
spp., or highly modified into palpacle-less oleocysts in P. vanhoeffeni. 
Also the female gonophores of the pyrostephids contain more than 
one ovum, while those of Erenna, where known, are monovan. 
These differences are here considered to be sufficient differences to 
warrant the establishment of a new family for the Erenna and closely 
related species described herein. The exact status of the genus 
Marrus remains uncertain, as Pugh (1999) discussed. 

Erenna Bedot, 1904 

Erenna Bedot, 1904: 10-14. 

Diagnosis. Nectophores dorso-ventrally flattened with tapering 
axial wings; apico- and infra-lateral ridges respectively form upper 
and lower margins of lateral surface, with short, perpendicular, 
vertical lateral ridge connecting them. Lateral radial canals straight, 
thickened on apico-lateral margins of nectosac; with or without 
additional small protuberances, spikes, or 'horn' canals. Bracts of 
two types, both with patches of epidermal cells, including 
nematocysts, on dorsal swelling at distal extremity. Tentillum large, 
with hypertrophied, uncoiled cnidoband, and rigid terminal process 
devoid of nematocysts. Gastrozooid with large swollen basigaster, 
but no obvious pedicle. 

Erenna richardi Bedot, 1904 

Erenna richardi Bedot, 1904: 10-14, PI. II, figs. 1-12 
Erenna bedoti Lens & van Riemsdijk, 1908: 66-69; Margulis, 1977: 
148-151, 1990: 138-142. 

Material examined. The description is based largely on a speci- 
men collected during Johnson-Sea-Link (JSL) II Dive 1456 (2 xi 
1987; 24° 00.9'N 82° 15.7'W; depth 871 m). In addition, parts of 
another large specimen collected during JSL I Dive 2889 (19 xii 
1990; 26°22.3'N 78°46.2'W; depth 701 m) have been examined. 
These parts have been donated to The Natural History Museum, 
London, where they are registered as BMNH 2000.1819. Unfortu- 
nately, the remainder of the specimen, including the siphosome, has 
dried up. 

DIAGNOSIS. Nectophores large, flattened, with prominent apico-, 
infra- and vertical lateral ridges; plus at least two pairs of indistinct 
and incomplete laterals in basal half; apico-laterals divide close to 
ostium. Thrust block large with, in mature nectophores, two small 
digitate protuberances on ventral surface. Radial canals black 
pigmented; lateral ones with thickened walls in region of lateral 
margin of nectosac; with small protuberances or spikes. Gastrozooids 
black pigmented, particularly in greatly expanded basigaster, with 
two prominent lateral lobes. Tentillum with hypertrophied cnidoband, 
and long rigid distal terminal process with a diverticular canal and a 
pair of 'ocelli' close to its end. 

Description. An image (Fig. 1 ) taken from a video of the in situ 
JSL II Dive 1 456 specimen shows the biserially arranged nectophores 
and the contracted siphosome. 

PNEUMATOPHORE. Pneumatophore ovoid, measuring 7 by 4 mm. 
Margulis (1977) noted the presence of eight vertical septa on her 
specimen, but these were not visible on the present material. 

Nectophore. (Figs 2 & 3). About 45 nectophores, at various 



ERENNA REVISION 



171 




Fig. 1 Erenna richardi. Image from in situ video of JSL II Dive 1456 
specimen; approximately 70-80 cm in length. 

states of development, and several nectophoral buds, remained with 
the JSL II 1456 specimen. They were flattened, and measured up to 
32 mm in length, 33 mm in width and 10 mm in height. The large 
axial wings tapered toward their apices (Fig. 2 A aw). Mature 
nectophore had relatively large thrust block (Fig. 2C tb) with a broad 
U-shaped indentation apically. On its ventral surface there were two 
small conical protuberances (Fig. 2C cp). However, on the younger 
nectophores, the thrust block was small and had no protuberances 
(Fig. 3). 

The prominent main ridge system consisted of pairs of apico- 
(Fig. 2A ral) and infra-laterals (Fig. 2B ril), which united close to 
the apex of each axial wing; and a pair of vertical laterals (Fig. 2B 
rvl) that connected the apico-laterals with the infra-laterals; al- 
though in some nectophores the junction with the latter was weak 



and difficult to discern. Apico-laterals branched close to the ostium, 
with each branch reaching the latter. In addition to these ridges at 
least two pairs of incomplete lateral ridges passed obliquely down 
the lateral facet in the basal half of the nectophore. Often these were 
difficult to discern without staining, but were also described by 
Margulis (1977), who noted 2-3 pairs of them. 

Ostial opening basal with no obvious mouth plate. On each side of 
the ostium there were prominent lateral protuberances on which lay 
strips of distinctive epidermal cells. A much smaller triangular patch 
also was present dorsally. In addition, there was a small patch of 
such cells on each side of the nectophore, at about the mid-height of, 
and basal to, the vertical lateral ridges. All these patches of epider- 
mal cells are believed to be sites of bioluminescence. 

T-shaped nectosac (Fig. 2B n) with a distinct muscle-free area 
across the whole of its apical region. Pallia! canal (Fig. 2A pa) was 
relatively short, running from the base of the thrust block over onto 
the ventral surface and ending just beyond the point of origin of the 
pedicular canal (Fig. 2B ped). On the nectosac the pedicular canal 
typically gave rise to all four radial canals, although occasionally 
there was a slight asymmetry in the arrangement. All four radial 
canals were straight. Laterals pass out, through the muscle-free 
zone, toward the lateral margins of the nectosac. Typically, before 
reaching the latter, they became thickened and could have small 
protuberances, or spikes extending up from them. These thickenings 
were particularly prominent on the youngest nectophores. All the 
canals had brown, but originally black, pigment in their gastrodermal 
walls. 

SlPHOSOME. As the in situ video (see Fig. 1) showed, the sipho- 
some was tightly contracted, and possibly, as in Agalma okeni 
Eschscholtz, 1825 and Frillagalma vityazi Daniel, 1966 (see Pugh, 
1998), this was its permanent state. 

Bract. (Fig. 4). Two types of bract were present; the first long, up 
to 50 mm, and narrow (Fig. 4A); the second shorter and broader (Fig. 
4B, C). Both possessed a pair of prominent, lateral cusps. These 




Fig. 2 Erenna richardi. A. upper and B. lower views of mature nectophore. C. detail of folded back thrust block. Scale bar 5 mm. aw axial wing; cp 
conical protuberance: n nectosac; pa pallial canal; ped pedicular canal; ral, ril, rvl apico-, infra- and vertical lateral ridges; tb thrust block. 



172 



P.R.PUGH 




Fig. 3 A. upper and B. lower views of young nectophore. Scale bar 5 mm. 




Fig. 4 Erenna richardi. A. first and B., C. second types of bract. D. detail of distal end of a bract. A-C. scale 2 mm; D. scale 0.2 mm. 



were situated in the distal half of the second type of bract, but in the 
first type they were positioned at about 13-14 mm from the proximal 
end of the bract, whatever the length of the latter. Thus in the longest 
bracts they were situated at about one quarter to one third its length; 
whereas in smaller ones they could be positioned in the distal half of 
the bract. Longer bracts of the first type had a second pair of lateral 
cusps close to the distal end. For both types, the bracteal canal 
originated, proximally. on the dorsal surface of the bract. For most of 
its course it remained in close contact with the ventral wall of the 
bract and there were striated bands of cells on each side of it. Close 
io the distal end of the bract it penetrated through the mesogloea to 
terminate below a small cup-shaped clump of large epidermal cells 
sunk into the dorsal surface at the tip of the bract (Fig. 4D). Some of 
these cells in this clump probably produced bioluminescence; others 
were nematocysts measuring c. 68 x 24 urn. No patches of such 
epidermal cells were noted elsewhere on the bract. 

GASTROZOODD. (Fig. 5 A, B). Large gastro/ooids contained dark 
brown (black in life) pigment. The proboscis region often was 



widely open and folded back on itself exposing a mass of villi. 
Enormous basigaster with two large lateral lobes, and also ex- 
panded, to a lesser extent, on the side opposite to where the tentacle 
was attached. No obvious pedicle. 

Tentacle and tentillum. (Figs 6, 7). Bedot's (1904) original 
description and illustrations of the tentacle and tentilla were detailed 
and accurate, and need little elaboration. The annulated tentacle had 
a muscular lamella running down one side, with the tentilla attached 
on the opposite side, at the internodes. 

Each tentillum consisted of a pedicle (Fig. lp), a cnidoband (Fig. 
7c) and a terminal process (Fig. Itp). The largest tentilla (Fig. 6) 
have a pedicle of up to 4-5 mm; longer than Bedot described. 
However, no doubt its length can be varied in life, and photographs 
of the specimen before preservation showed the pedicle to be highly 
contracted. The cnidoband measured up to c. 15 mm long and was 
laterally compressed. It consisted of the cnidoband proper, where 
the nematocysts are attached, and what Totton (1965) called, in the 
case of Pyrostephos vanhoeffeni Moser, 1925, the saccus (Fig. 7.?). 



ERENNA REVISION 



173 




Fig. 5 Erenna richardi. A. and B. Gastrozooids; C. and D. palpons; E. 
Immature gonodendron. Scale 2 mm. 

The bulk of the saccus was made up of a thick layer of transparent 
gastrodermal cells which formed a characteristic reticulate pattern 
(Fig. 6), through which the narrow, dark (black in life) pigmented 
gastrovascular canal (Fig. Igvc) passed. In many preserved tentilla 
this canal was damaged. Figure 6 shows an undischarged and a 
discharged tentillum. 

The numerous nematocysts formed a dense, darkly pigmented 
cnidoband, the sides of which undulated irregularly. On the mature 
tentillum the distal end of the cnidoband hung free from the main 
body. Three types of nematocysts were present. On the sides there 
were rows of larger ones, probably homotrichous anisorhizas, meas- 
uring c. 165 x 32 pm. Between these were numerous smaller 
nematocysts of two types, one measured c. 43 x 15 pm, the other c. 
27 x 20 pm. No discharged nematocysts of these types were found, 
but probably they were both atrichous haplonemes, as Margulis 
(1977) suggested, although she noted only one type. 

The rigid terminal process (Fig. 6) was up to c 10 mm in length 
and devoid of nematocysts . The gastrovascular canal passed through 
it, ending close to its distal extremity. Just proximal to the end of the 
canal it branched off a diverticular canal (Fig. Idc) that passed back 
through the terminal process to end below the distal part of the 
cnidoband. As Bedot (1904) noted there was a band of longitudinal 
musculature on each side of the terminal process. Overlying the 
lateral sides of the diverticular canal, close to its point of origin, 
there was a pair of brownish-white oval structures (Fig. 7o) com- 
prised of distinctive epidermal cells. Lens & van Riemsdijk (1908) 
aptly called them 'ocelli', as is discussed below. 

As noted above it was the distal end of the cnidoband that hung 
free from the main body of the mature tentillum. However, on the 
young, developing tentillum (Fig. 7A), the cnidoband formed a 
triangular process that was slightly undercut on its proximal surface; 



with the pedicle and terminal process being only slightly developed. 
The axial and diverticular canals were prominent, and the latter 
opened into the saccus of the cnidoband. With further development, 
the saccus became closed off and began to fill with gastrodermal 
cells (Fig. 7B). The cnidoband remained undercut proximally. With 
further elongation of the tentillum (Fig. 7C, D), the proximal part of 
the cnidoband began to fuse with the saccus, and the 'ocelli' on the 
terminal process were developed, while the canal system narrowed. 
Finally, the distal end of the cnidoband became detached from the 
saccus. 

PALPON. (Fig. 5C, D). Up to 15 mm in length, with a palpacle 
attached at the base of the pedicle. Brown (black in life) pigment 
throughout. Pedicle with reticulate pattern of cells. Main stomach 
region with an irregular pattern of patches. Distally these were 
concentrated to form 12-14 vertical stripes, with denser pigmenta- 
tion, surrounding the base of the proboscis with its terminal opening. 
No obvious nematocysts present on the palpacle. 

GONODENDRA. (Fig. 5E). Only female gonodendra were found on 
the JSL II 1456 specimen. Mature female gonodendra were com- 
prised of small, tightly packed bunches of c. 20-25 gonophores, with 
a milky brown coloration, connected to a relatively short stalk. Each 
gonophore measured c. 0.45 mm in diameter and contained a single 




Fig. 6 Erenna richardi. Photographs of tentilla (c. 25-30 mm in length) 
before (A.) and after (B.) discharge of nematocysts. 



174 



P.R.PUGH 




gvc 



Fig. 7 Erenna richardi. A-D. Early stages in the development of a 
tentillum. Scale 0.5 mm. c cnidoband; dc diverticular canal; gvc 
gastrovascular canal; o 'ocellus';/; pedicle; tp terminal process. 

egg. The immature gonodendra (Fig. 5E) were more darkly pigmented 
and had a relatively long stalk which typically bifurcated close to its 
apex, with the gonophores being developed on the branches. One of 
these branches could be denuded and could be mistaken for a 
gonopalpon. Occasionally small gonophores were budded off 
approximately half way up the stalk. No gonophores were found 
with the JSL I Dive 2889. However, two specimens from the 
Discovery collections also bore only female gonodendra. 

DISTRIBUTION. Of the four specimens of Erenna richardi col- 
lected by the JSL submersibles, one came from the vicinity of the 
Dry Tortugas, between Florida and Cuba; two from The Bahamas; 
and one from c. 27°N 85°W in the Gulf of Mexico (Rebeca Gasca - 
personal communication). The species also has been found occasion- 
ally in recent Discovery collections in the North Atlantic, mainly 
south of 35°N and at depths greater than 1000 m. Much of the 
material is in too poor a condition to identify to species, although it 
seems likely that the presence of black pigment in the radial canals 
of the nectophore is specific to E. richardi. However, one large 
specimen, comprising 34 nectophores, over 400 bracts and several 
stem pieces, from Discovery St. 8599 (c. 8°40'N 23°14'W; depth 
300O-3500m). clearly is E. richardi: as is a specimen from St. 
I0157#3(31°3.6'N 13°W; 650-1000 m). In addition some damaged 
nectophores that can be referred to this species have been collected 
by four hauls fished C.300 m above the bottom in the proximity of the 
Rainbow' and 'Lucky Strike' vents on the Mid-Atlantic Ridge (c. 



37-38°N, 32-33°W; depth range 1793-2613 m). Better preserved 
material has also been collected recently in the vicinity of the East 
Pacific Rise (c. 12°50'N, 104°W) at depths of about 2500 m. The 
siphosome of one of these specimens bore numerous female 
gonodendra. 

Leloup ( 1 980) listed the scant published data on the geographical 
distribution of Erenna richardi, including the conspecific E. hedoti. 
The material described by Bedot (1904), Lens and van Riemsdijk 
(1908), Bigelow (1911), Moser (1925) and Leloup (1936, 1980) 
probably can be referred to E. richardi; but this is not the case for all 
of Totton's (1965) material. His figure 38, which is said to be a 
reconstruction, shows a nectophore with short 'horn' canals arising 
from the lateral radial canals on the nectosac. Re-examination of the 
single nectophore, in the NHM collections, shows that incomplete 
lateral ridges are also present and so it is referred to E. richardi. 
However, the label with it states that the Discovery station at which 
it was collected is St. 4255 not, as Totton stated, St. 2061. The 
reverse is true for the specimen figured in figure 39. Since figure 39 
looks more like a 'reconstruction' than figure 38, it is presumed that 
the legends to the two figures were accidentally transposed. Totton 
(1965) mentioned a larger specimen from Discovery St. 4230, but 
this was not found. His other material from La Jolla, California, 
supplied by Dr Ahlstrom, and the Beebe collections from Bermuda, 
together with some bracts from Discovery St. 4253, which he did not 
mention, probably can be referred to E. richardi, but are in poor 
condition. The Discovery St. 72061 material, is referred to another 
species, described below. 

Of the other records not included by Leloup ( 1 980) or published 
more recently, no description was given by Alvarifio ( 1 969, 1 980) or 
Musayeva (1976) and so their true identity remains in doubt; par- 
ticularly since Alvarifio (1981) described a specimen that is probably 
not E. richardi. However, as with the description of Daniel (1985), 
it is difficult to decide whether or not she was largely copying the 
description and figures of Totton (1965). The two specimens 
described or mentioned by Margulis (1969, 1977, 1990) can be 
referred to E. richardi, although in the last paper the specimen was 
described under the name E. bedoti. This will be discussed in more 
detail below. Finally, the record given by Pugh (1975) is based on E. 
richardi. 

Behaviour. The JSL II 1456 specimen was briefly observed on 
board the mother ship after collection. It was noted that the terminal 
processes of the tentilla were kept rigid and were rapidly vibrated. 
Meanwhile, the cnidoband might become bent into a U-shape. It has 
been suggested (Pugh, 1989) that vibration of the terminal process 
might be an example of aggressive mimicry of a swimming 
chaetognath. The two 'ocelli' might then be a representation of 
chaetognath's gonads. Equally, the vibration of the terminal process 
might be mimicking the swimming behaviour of a larval fish; the 
two 'ocelli' then representing eyes. 

As Totton ( 1 965, p.76) noted, 'What is so characteristic of Erenna 
[richardi] is the hypertrophy of the cnidoband, which must be a very 
formidable stinging apparatus'. This has certainly proved to be the 
case, as was demonstrated when a colleague was painfully stung 
when he inadvertently came into contact with a tentacle of one of the 
specimens collected by the JSL submersible. 

Remarks. Lens and van Riemsdijk's ( 1 908) description of Erenna 
hedoti was based on two fragments of stem; one being the nectosome, 
with an apical pneumatophore, and the other a small part of the 
siphosome. However, they could only compare the tentacles and 
tentilla with those described by Bedot's ( 1904) for E. richardi. For 
the tentacle they stated (p. 68) that it 'reminds one exactly of the 
tentacle described by Bedot'; and (p. 69) that'the most mature 



ERENNA REVISION 



175 



tentillum . . . reminds one at once of the tentilla described by 
BEDOT'. They concluded (p. 69) that 'there exists undeniably the 
closest relationship between the tentacles and tentilla of Erenna 
Richardi and Erenna Bedoti.' So why did they separate them? Their 
earlier logic (p. 66) 'We therefore called this only specimen Erenna, 
using a new species denomination "Bedoti" as of course we cannot 
decide whether the tentacles described by BEDOT belonged to a 
specimen entirely identical with ours' seems very obscure. 

Lens and van Riemsdijk suggested a possible difference between 
the tentilla of their specimen and that of Bedot. They believed that, 
in the largest tentillum, the free end of the cnidoband was proximal 
('basal') while in Erenna richardi it was distal (although they 
referred to the latter as 'proximal'). They also noted the presence of 
a small black spot in the distal region of the cnidoband, which they 
suggested would become the 'ocelli' that Bedot described. How- 
ever, the 'ocelli' off. richardi are positioned toward the end of the 
terminal process, even in the developing tentilla (see Fig. 6C, D). 
Have, then, Lens and van Riemsdijk misinterpreted the structure of 
their tentilla? Close examination of their illustration (Plate XI, fig. 
89) suggests that this is probable. It is suggested that what they call 
the pedicle is, in actuality, a very deformed terminal process; while 
the 'apical part' (terminal process) is the pedicle. Then the free end 
of the cnidoband is distal, as it is in E. richardi. For this to be so, the 
spot, which lay close to the proximal end of the cnidoband, and 
which they thought was equivalent to the 'ocellus' of E. richardi, 
must be considered an artefact, and that the true 'ocelli' have been 
destroyed. This is borne out by Lens and van Riemsdijk ( 1 908, p. 69) 
statement that 'Microscopical sections have been made but the 
material is unfortunately absolutely insufficient, the different layers 
being all destroyed'. It appears that all the largest tentilla were 
sectioned as none are now present with the type material. However, 
some developing tentilla are still present with the holotype and these 
conform exactly with those of E. richardi. Thus, there does not 
appear to be any concrete evidence to separate specifically Lens and 
van Riemsdijk's material from that of Bedot. 

This conclusion was reached by Totton (1965). who considered 
Erenna bedoti to be conspecific with E. richardi. However, 
Margulis (1977, 1990) resurrected the debate, when she described 
another specimen of E. richardi Margulis (1997). She correctly 
noted that, on the nectophores, there were two digitate processes 
on the ventral side of the thrust block, and 2-3 extra ridges on the 
lower lateral facet. On the siphosome she found, as indeed had 
Lens and van Riemsdijk (1908), the presence of peculiar muscular 
outgrowths, but did not know their function. They are, undoubt- 
edly, the remains of the muscular lamellae to which bracts were 
once attached. One curious feature she described was that one 
detached tentacle arose from a black-pigmented formation that 
had a spherical dilation on each side. This was, of course, the 
basigaster of the gastrozooid, but she failed to appreciate this. The 
youngest tentillum had a digitate outgrowth in the region that was 
to become the cnidoband; the oldest were as Bedot (1904) 
described them. Margulis (1977) then made a brief comparison 
with Lens and van Riemsdijk's (1908) description of E. bedoti. 
The only differences she noted were that the tentacles of E. bedoti 
lacked the basal outgrowths (i.e. the basigaster), and that their 
young tentilla did not have a digitate outgrowth. 

These points were addressed further by Margulis (1990), when 
she described fragments of another specimen that she referred to 
Erenna bedoti. The key features by which she distinguished this 
specimen from E. richardi were that:- 

a) the thrust block on the nectophore was smaller, with its distal 
margins stretched into thread-like outgrowths; and that the two 



processes on its ventral surface were digitate or papillose, but not 
lamellate as in Erenna richardi; 

b) there were marked differences in the structure of the gastrozooid, 

and; 

c) the young tentilla of Erenna bedoti had an oval outgrowth, while 

in E. richardi it was finger-shaped. 

As has been shown above, the size of both the thrust block and the 
digitate processes varies with the size of the nectophore of Erenna 
richardi, which, as will be seen, is the only erennid species to have 
such processes. Thus, judging from her illustrations, the damaged 
condition of Margulis's material is the most likely explanation of the 
differences she noted. It should be noted, moreover, that black 
pigment was present in Margulis's material. The differences in the 
gastrozooids mainly concern the basigaster. Margulis now recog- 
nised that, in Erenna richardi, this was the large structure, with two 
large rounded lobes, that she had found at the proximal end of the 
tentacle. However, such a structure was not found on the larger 
gastrozooids of her latest specimen; although it was clearly large and 
inflated on the younger ones. This would be a reasonable difference 
if Margulis had not given the impression that the basigaster had been 
destroyed, by describing how its outer coating began to shed until it 
was completely absent. Personal experience has shown that it is, 
indeed, easy to destroy the epidermal layers of the basigaster. 
Finally, the differences between the young tentilla are considered to 
be mere reflections of their state of growth. The important similari- 
ties between Margulis's (1990) material and E. richardi are that in 
both there is a pair of protuberances on the ventral side of the thrust 
block of the nectophore, and that black pigmentation is present. The 
former, and almost certainly the latter, of these are characteristics for 
only E. richardi and, thus, it seems inconceivable that E. bedoti is 
nothing more than conspecific with E. richardi. 

Erenna laciniata sp.nov. 

Holotype. The specimen from JSL II Dive 1454 is designated 
holotype, and has been donated to the Natural History Museum, 
London where it is registered as BMNH 2000.1821. 

Material examined. The description is based on two specimens 
collected by the JSL II submersible during dives 1454 (30 viii 1987; 
24°30.8'N 83° 45.2'W; depth 81 1 m) and 1688 ( 1 1 xl988; 26°23.5'N 
78°39.5'W; depth 853 m). 

Diagnosis. Nectophores large, dorso-ventrally flattened, with 
only basic ridge pattern; with weak division of apico-laterals close to 
ostium. Thrust block small, with U-shaped median indentation and 
ventral flaps, but no conical protuberances. Lateral radial canals 
only slight thickened at apico-lateral corners of nectosac. Bracts of 
two types, with lateral flap, more extensive in one type than the 
other. Tentilla characteristic, with terminal process arising close to 
base of cnidoband and bearing two distal 'ocelli'. 

Description 

PNEUMATOPHORE. The pneumatophore measured c. 2.9 x 1 .9 mm; 
gas expansion having ruptured its base. There were no obvious 
striations or pigment. 

NECTOSOME. Each specimen had 50-60 nectophores, which, in 
life, were arranged biserially. 

Nectophore. (Figs 8, 9). Flattened, up to 25 mm in length, 29 mm 
in width and c. 5 mm in height. Large tapering axial wings. On the 
mature nectophores (Fig. 8) the thrust block was relatively small and 
divided into two parts by a median U-shaped indentation. 



176 



P.R.PUGH 




Fig. 8 Erenna laciniata. A. upper and B, lower views of mature nectophore. Scale 5 mm. 



On each side there was a thickened flap that lay ventrally and was 
directed toward the mid-line. On smaller, younger nectophores 
i Fiy. 9) the thrust block consisted of two small protuberances sepa- 
rated by a U-shaped indentation. 

The main ridge system was well delineated, especially in the 
younger nectophores (Fig. 9). It consisted of pairs of infra- and apico- 
lateral ridges, which united close to the lateral apices of the axial 
wings; and a short pair of vertical laterals connecting them (Fig. 8), 
although, in younger nectophores, the junction with the apico-laterals 
was indistinct (Fig. 9). The infra-laterals ended, basally, on either side 
of the ostium. The apico-laterals remain prominent until just above 
ostial level. In younger nectophores, close to the ostium, the apico- 
laterals gave rise to three very vague, but broad branches (Fig. 9A). 
However, in mature nectophores only after staining could two very 
vague ridges be discerned (Fig. 8A). These would be virtually 
impossible to see in damaged or poorly preserved material. 

No obvious mouth plate, and ostial opening basal. Prominent 
protuberances on each side of the ostium which bore strips of 
distinctive epidermal cells, and another triangular patch in mid-line 
on dorsal side of the ostium. In addition, a relatively large patch of 
such cells was present on each side of the nectophore, at about the 
mid-height of, and immediately basal to, the vertical lateral ridges 
(Figs 8A, 9A). All these patches are believed to be sites of bio- 
luminescence. 

T-shaped nectosac with a distinct muscle-free zone at its apex. 
Short pallial canal, originating at the base of the thrust block, with a 
long pedicular canal, which on reaching the nectosac gave rise to the 
four, straight, radial canals. In the younger nectophores there were 
obvious signs of thickening of the lateral radial canals in the apico- 




lateral region of the nectosac, but these were difficult to discern in 
the mature ones. No small protuberances, or 'horn' canals, were 
present. In the JSL material the radial canals had no obvious 
pigmentation, although in other nectophores, which are tentatively 
referred to this species, they could have an orange hue. 

SlPHOSOME. On collection the siphosome of both specimens was 
tightly contracted. The gastrozooids and the terminal processes to 
the tentilla were lightish brown in colour; while the palpons were 
suffused with brown pigment. 

Bract. (Fig. 10). Over 1 000 bracts, oftwo types, were found with 




Fig. 9 Erenna laciniata. A. upper and B. lower views of immature 
nectophore. Scale 2 mm. 



Fig. 10 Erenna laciniata. Bracts. A., B. dorsal views of first type; C, D. 
dorsal views of second type. Scale 5 mm. E. Immature bract. Scale 1 mm. 



ERENNA REVISION 



177 



the JSL 1688 specimen, occurring as enantiomorphic (mirror-im- 
age) pairs. The first type of bract (Fig. 10A, B), which measured up 
to 25 mm in length, was deeply incised on one side, at about its mid- 
length where a large flap was formed stretching across almost to the 
mid-line. On the other side, slightly proximal to this level, a small 
cusp was present. Toward the distal end, there were two further 
lateral cusps, with the one on the same side as the flap tending to be 
more marked than the other. The distal end of the bract was pointed. 
Close to the tip, the dorsal surface was thickened into an oval patch 
of cells that surround and were interspersed with over 200 
nematocysts, measuring c. 63 x 25 urn. Another small patch of such 
cells was found on the dorsal surface of the bract, in the mid-line, 
and on a level with the lateral flap. That patch, at least, is believed to 
be a site of bioluminescence. Proximally, the bracteal canal origi- 
nated on the dorsal surface of the bract. It then curved over onto the 
ventral surface and continued distally in the mid-line. It appeared to 
be of variable thickness, but such variability was the result of 
variations in the thickness of the striated tissue that lay on each side 
of it. Close to the distal end of the bract the canal penetrated into the 
mesogloea and ended below the oval patch of epidermal cells. 

The second type of bract (Fig. IOC, D) was similar to the first, but 
tended to be shorter, up to 20 mm in length, and broader. The lateral 
flap, however, was much reduced, although still an obvious feature. 
Distal to this again there was an obvious lateral cusp; but on the other 
side the cusp was very small or absent altogether. The first type of 
bract was about three times more numerous than the second. Roughly, 
with over a thousand bracts and approximately 25 gastrozooids, 
there would seem to have been about 40 bracts per cormidium. Very 
young bracts (Fig. 10E) were roughly pyramidal in shape with the 
distinctive patch of epidermal cells, including nematocysts, fully 
developed. The bracteal canal was short and did not extend onto the 
dorsal surface; while beneath the distal patch of epidermal cells it 
formed an extensive cavity. 

GASTROZOOID. (Fig. 1 1 A, B). The gastrozooids measured up to c. 
1 5 mm in length. The proboscis region, which often was curled back 
over itself, bore some stripes of gastrodermal cells. The stomach 
region, externally, was featureless and had a brown colour. The 
basigaster was greatly expanded on all sides, except that to which 
the tentacle was attached, and there was no obvious pedicle. 

Tentacle and tentillum. (Fig. 12). Typically the annulated 
tentacle had a muscular lamella running down one side, with the 
tentilla attached on the other side, at the internodes. The tentilla were 
of an extraordinary design. Only in the very young tentilla (Fig. 12 - 
centre) was there any trace of a pedicle. In these the cnidoband was 
made up of a large saccus overlain by a horseshoe-shaped band of 
nematocysts. The terminal process actually arose from the base of the 
saccus and bore, towards its distal end, a pair of 'ocelli'. The 
gastrovascular canal penetrated through the terminal process, but no 
connection with the saccus of the cnidoband could be discerned. As 
the tentilla matured the cnidoband lengthened, with the terminal 
process still only attached close to its base. The saccus diminished in 
importance and distally the band of nematocyst occupied all but a 
narrow strip of the external surface of the cnidoband. Proximally, 
where the terminal process was attached, the band of nematocysts split 
into two parts on either side of the saccus. Three types of nematocysts 
were present. The large anisorhizas, which measured c. 1 28 x 27 urn, 
were arranged along the lateral margins of the cnidoband. Between 
them were numerous smaller nematocysts of two shapes, probably 
heteronemes, with the more cylindrical ones measuring c. 40 x 1 5 um 
and the more ovoid ones c. 32 x 18 um. Bands of musculature were 
present in the terminal process and extended distally to beneath the 
'ocelli'. In life this process was reddish-brown in colour. 





Fig. 11 Erenna laciniata. A., B. gastrozooids; C. palpon. Scale 2 mm. D. 
male gonophores. Scale 1 mm. 




Fig. 12 Erenna laciniata. Three mature tentilla (Scale 1 mm), with 
(centrally) an immature one (Scale 0.5 mm). 

Palpon. (Fig. 11C). Numerous palpons, up to c. 15 mm long, 
were present with the specimens. They were featureless thin-walled 
sacs filled, with a milky-white amorphous substance, although in 
life they were suffused with a brownish hue. The extent of the 
proboscis was variable, but typically, at its base, there were some 



178 



P.R.PUGH 




Fig. 13 Erenna laciniata. Photograph of part of siphosome showing male 
gonophores. 

short, brown-coloured stripes. No nematocysts were found on the 
palpacle. 

Gonodendra. (Figs 1 ID, 13). The JSL 1688 specimen was fe- 
male and the gonodendra were arranged in a very similar fashion to 
that described in Erenna richardi. The JSL 1454 specimen was 
male, with the individual gonophores apparently scattered randomly 
down the stem (Fig. 13). Each was a relatively large medusoid, 
whose manubrium progressively filled with spermatozoa until it 
occupied almost the entire subumbrella cavity (Fig. 1 ID) and had a 
milky-white colour. 

Distribution. The JSL II Dive 1454 specimen came from the 
region of the Dry Tortugas between Florida and Cuba, while the 
Dive 1688 specimen came from the region of The Bahamas. In 
addition, some damaged nectophores that can be referred to this 
species have been found in the Discovery collections from indi- 
vidual hauls on the equator at 22°W (805-900 m) and at 3°N 23°W 
(0-1000m). 

Behaviour. In life the terminal process of the tentillum was 
reddish-brown in colour. It was kept rigid and was vibrated rapidly 
presumably, as was suggested for Erenna richardi, as a form of 
aggressive mimicry. 

ETYMOLOGY. The specific name, being Latin for a 'flap', refers to 
the lateral flap-like process on the bracts. 

Erenna cornuta sp.nov. 

HOLOTYPE. The JSL II Dive 1451 specimen is designated holotype, 
and has been donated to the Natural History Museum, London 
where it is registered as BMNH 2000.1818. 

Material examined. The description is based on a single speci- 
men collected during JSL II Dive 145 1 at a depth of 896 m. (29 viii 
1987; 24°30.6'N 83°45.6'W). In addition, a few specimens that 
apparently can be referred to this species have been found in recent 
Discovery collections, but always in a poor state of preservation. 

DIAGNOSIS. Nectophores relatively less dorso-ventrally flattened, 
with only the basic ridge pattern; with apico-laterals not dividing 
close to ostium. Thrust block small, with no median indentation or 
conical protuberances. Lateral radial canals typically have 'horn' 
canals branching off at apico-latcral margins of ncctosac. Bracts 




Fig. 14 Erenna cornuta. A. upper and B. lower views of nectophore with 
'horn' canals; C. upper view of nectophore without 'horn' canals. D. 
upper view of young nectophore. Scale 2 mm. 

with weak transverse ridge. Tentilla of two characteristic types; one 
with a long pedicle and no terminal process, with nematocysts 
grouped into four circular processes; the other with shorter pedicle 
and a small vesicular terminal process; with nematocysts, on the 
long cnidoband, more or less separated into c. 17 bundles. 

Description 

PNEUMATOPHORE. Measured c. 1.5 x 1 mm, but its base has been 
ruptured by gas expansion. No obvious pigmentation. 

Nectophore. (Fig. 14). Thirteen large nectophores, plus two 
developing ones and some nectophoral buds, were found with the 
specimen. They measured up to 16 mm in length, 14 mm in width 
and 4 mm in depth. The ridge pattern comprised pairs of apico-, 
infra- and vertical laterals. The apico- and infra-laterals unite close 
to the apices of the axial wings. The infra- laterals extended basally 
to end below the ostium. The apico-laterals, on mature nectophores, 
were only prominent in the upper half of the nectophore and, unlike 
the two species described above, basal to the vertical lateral ridges, 
they curved in toward the mid line (Fig. 14A, C). They could be 
traced, usually only after staining, down further toward the ostium, 
but did not divide. On the youngest nectophore the apico-laterals 
rapidly approached and then overlapped each other, ending just 
above ostial level (Fig. 14D). 

No obvious mouth plate and the ostium opened basally. On each 
side of the ostium there were prominent lateral processes with 
obvious strips of epidermal cells; with a smaller triangular patch of 
cells found dorsally. In addition, there was a pair of small patches of 
cells on either side of the nectophore; above the mid-height of and 
basal to the vertical lateral ridges. All these patches were believed to 
be sites of bioluminescence. 

T-shaped nectosac with a distinct apical muscle-free area, par- 
ticularly on its ventral side. Pallial canal short, extending from the 
base of the thrust block over on to the ventral surface of the 
nectophore, where it gave rise to the pedicular canal. On the 
nectosac the latter immediately divided to form the four straight 
radial canals. No pigmentation was noted in any canals. The lateral 



ERENNA REVISION 



179 




Fig. 15 Erenna cornuta. A. Three bracts of first type; B. two bracts of 
second type. Scale 2 mm. 

radial canals passed through the middle of the muscle-free zone to 
reach the apex of the nectosac just short of its lateral margins. At 
that point the narrow 'horn' canals may or may not be given off. 
When present these 'horn' canals extended up for a variable dis- 
tance, toward the apex of the nectophores, typically terminating 
on a level with the central thrust block. Eight nectophores (Fig. 
14A, B) had well-developed 'horns', as did the two immature 
ones (Fig. 14D). In three others the 'horn' canals were ill-defined 
and short; while in the remaining two (Fig. 14C) there was no 
trace of them whatsoever. 

The mature nectophores varied slightly in shape apparently in 
association with the extent of the 'horn' canals. The tapering axial 
wings were more extensive on those with well developed 'horn' 
canals; while the central thrust block typically was slightly smaller 
than those without 'horn' canals. The ones with ill-defined 'horn' 
canals tended to be intermediate. Of the 23 damaged nectophores 
collected at Discovery St. 7856#54 the 'horn' canals were prominent 
in all but the two smallest, but mature ones. The others were larger 
than the JSL II Dive 1451 ones, measuring c. 20 mm in length and 
width, and had denser musculature on the nectosac. The 'horn' 
canals also were more extensive, and there were traces of orange- 





Fig. 16 Erenna cornuta. A. gastrozooid, and B. palpon. Scale 1 mm. 



brown pigment in the basal parts of the radial canals and, particu- 
larly, the ostial ring canal. 

Bracts. (Fig. 15). Two types of bract were present; the first (Fig. 
15 A), and considerably more numerous, being longer, up to 16 mm 
in length, and narrower than the second (Fig. 1 5B ), which was up to 
1 mm in length. Both types possessed a pair of lateral cusps. In the 
first type, as was the case with Erenna richardi, these seemed to lie 
at a fixed distance from the proximal end of the bract, and were 
asymmetrically disposed. Along the axis between these two cusps, 
on the dorsal surface of the bract, there was a rounded transverse 
ridge or process, that marked a change in the thickness of the bract, 
which was thinner distally. This ridge did not connect with the cusps, 
and its extent and distinctiveness was variable. Just distal to the 
ridge, in the mid-line, there was a small patch of cells; although often 
these has been abraded away. 

On the second type of bract the lateral cusps were positioned, 
almost symmetrically, at about the mid-length of the bract. Just 
distal to these cusps there was a more or less pronounced cross-ridge 
which again demarcated a change in the thickness of the bract. 
Again a small patch of cells was situated distal to this ridge, in the 
mid-line. On the distal half of both types of bract, there could be an 
additional lateral protuberance of variable shape. At the distal tip of 
both types the dorsal surface was raised up in the mid-line to form an 
elongate or elliptical process on which were found a concentration 
of small epidermal cells, with brownish-red pigment. Centrally, 
these cells included some nematocysts, which measured c. 68 x 
32 um. 

The bracteal canal originated, proximally, slightly over on the 
dorsal surface of the bract. It passed down the middle of the bract in 
close contact with its ventral wall, with striated bands of cells lying 
on either side of it, indicating where the muscular lamella was 
attached. At some distance from the distal end of the bract, in 
comparison with Erenna richardi, the canal penetrated into the 
mesogloea and curved up to end beneath the proximal part of the 
concentration of cells on the dorsal surface. 

Gastrozooid. (Fig. 16A). Only three well-developed gastro- 
zooids remained with the specimen. The largest was 6 mm in length. 
The basigaster formed a horseshoe-shaped, laterally expanded struc- 
ture around the base of the gastrozooid, with the tentacle attached in 
the open zone. It was a light brown colour. No obvious pedicle. The 
expanded stomach was externally featureless and had a dark brown 
colour. The proboscis region, which was about the same length as 
the stomach, had distinct stripes. 

Tentacle and tentillum. (Figs 17, 1 8). The tentacle was annu- 
lated, with a muscular lamella running down one side, and the 
tentilla attached, on the opposite side, at the internodes. There were 
two types of tentilla, both of which were found attached to the same 
tentacle. Early on in the development of the first type (Fig. 17 A), the 
tentillum consisted of a long pedicle and a minute cnidoband devoid 
of nematocysts. The gastrovascular canal was seen, at the end of the 
pedicle, to turn back and continue down to open into the cavity of the 
saccus of the cnidoband. With further development the connection 
with the saccus cavity was closed, and the cavity filled with 
gastrodermal cells. A remnant of the diverticular canal seemed to 
persist, passing through a relatively dense band of gastrodermal 
cells. In the mature tentillum (Fig. 17B), the proximal part of the 
pedicle was expanded, typically tapering towards its base. The 
gastrovascular canal could occupy most of the interior, or could 
remain as a narrow tube, which became twisted and folded up on one 
side of the pedicle. Distally the pedicle was narrower, with the canal 
often having a zigzag appearance, probably indicating some 



ISO 



P.R.PUGH 





Fig. 17 Erenna comma. A. young tentillum of first type. Scale 0.5 mm. 
B. Two views of mature tentillum of first type. Scale 1 mm. 

contraction in the length of the pedicle itself. The canal penetrated 
through the saccus of the cnidoband to its tip. The saccus was largely 
filled by large, vacuolate gastrodermal cells that formed a reticulate 
pattern. The nematocysts were restricted to four circular swellings; 
two lateral, one proximal, one distal. Three types of nematocysts 
were present and arranged so that the larger anisorhizas, measuring 
c. 130 x 35 urn. surrounded the smaller (?) haplonemes; the more 
cylindrical ones measuring 32 x 1 1 urn, and the more ovoid ones 41 
x 14 urn. 

The youngest of the other type of tentillum (Fig. 1 8A) comprised 
a short, thickened pedicle, a long cnidoband, and a short terminal 
process, which had a circular spot of distinctive epidermal cells on 
one side. The broad gastrovascular canal was present throughout 
and. in the terminal process, it appeared to bend back and continue, 
for a short distance, toward the saccus of the cnidoband. However, 
even in the smallest tentilla examined, the saccus had already been 
occluded by gastrodermal cells. The nematocysts had begun to 
accumulate into an undulating series of connected swellings on one 
side of the saccus. 

Only a few mature tentilla (Fig. 18B) of the second type were 
found with the specimen. In these the pedicle remained short and 
broad and was largely filled by the gastrovascular canal. The 
cnidoband had increased greatly in length and was largely filled by 




Fig. IS Erenna cornuta. A. Young tentillum of second type. Scale 0.5 
mm. B. Mature tentillum of second type. Scale 1 mm. 



large, vacuolated gastrodermal cells. The narrow gastrovascular 
canal passed through its middle and opened into the cavity of the 
small, spherical, thin-walled terminal process. The patch of cells 
persisted on one side of the latter. No diverticular branch of the main 
canal could be discerned. On the cnidoband the rounded swellings 
containing the nematocysts became more or less separated one from 
another. On the two best preserved mature tentilla there were 17 of 
these patches. The nematocysts were of the same type and size as on 
the other type of tentillum. 

Palpon. (Fig. 16B). The globular, thin-walled palpons meas- 
ured up to c. 4 mm in length. There was a short, narrow proboscis, 
at the base of which was a ring of pigmented gastrodermal cells, 
often organised into distinct spots. Other concentrations of 
gastrodermal cells sometimes were visible, particularly on the 
distal part of the stomach region. The base of the stomach region 
was almost surrounded by a small, loosely attached, horseshoe- 
shaped region of large, vacuolated epidermal cells, with the 
palpacle being attached in the open region. No nematocysts were 
found on the palpacle. 

GONODENDRON. No gonodendra were found on the small piece of 
siphosome that remained with the specimen. 

DISTRIBUTION. The type specimen came from the region of the 
Dry Tortugas, between Florida and Cuba. Nectophores with 'horn' 
canals, which presumably can be referred to this species, have been 
collected at four recent Discovery stations. Two of these were at c. 
30°N 23°W at depths of 1250-1500 m and 1500-2000m, and the 
other two from off Bermuda (c. 31°45'N 63°45'W) at depths of 
1250- 1500m. 

Etymology. The specific name, meaning 'horned' in Latin, re- 
fers to the 'horn' canals present in most of the nectophores. 

Parerenna gen. nov. 

Diagnosis. Nectophores not dorso-ventrally compressed; with 
muscle-free zone on nectosac mainly on lower surface adaxially. 
Vertical lateral and incomplete infra-lateral ridges very indistinct; 
the latter not forming the lower margin of lateral surface. Apico- 
laterals peter out well above ostial level. Gastrozooid with minute 
basigaster. Tentillum with long pedicle; with cnidoband extending 
beyond terminal process, which has a small spherical distal swell- 
ing. 

Monotypic genus to accommodate Parerenna emilyae sp.nov. 

Parerenna emilyae sp.nov. 

HOLOTYPE. The specimen from JSL 1 Dive 2886 is designated 
holotype, and has been donated to the Natural History Museum, 
London where it is registered as BMNH 2000.1820. 

Material examined. A single specimen collected during JSL I 
Dive 2886 (18 xii 1990; 26°31.8'N, 78°05.6'W; depth 823 m). 
Before preservation in 5% buffered formalin, the bioluminescence 
of the specimen was studied which, unfortunately, resulted in the 
loss of some parts. 

Diagnosis. As for genus. 

Description 

Pneumatophore. The base of the pneumatophore has exploded 
due to the expansion of the gas contents while bringing the specimen 
to the surface. Pneumatosaccus spherical, c. 1 mm in diameter, with 
a small cap of cells, which may have been pigmented in life. 



ERENNA REVISION 

A 



181 




Fig. 19 Parerenna emilyae. Nectophores. A., B. upper, C. lower, and D. 
lateral views. Scale 2 mm. 

Nectophore. (Fig. 19). Eight fully developed and two developing 
nectophores were found with the specimen; plus a few nectophoral 
buds at the apex of the highly contracted nectosome. Mature 
nectophores were not dorso-ventrally flattened and measured up to 
13 mm in length and width and 6 mm in height, and had well 
developed tapering axial wings. The central thrust block was broad, 
but of little height. The only obvious ridges were the apico-laterals 
(Fig. 19A, B), running down from the apices of the axial wings 
toward the ostium, but petering out well above that level. Only by 
staining were the pairs of complete vertical lateral and incomplete 
infra-lateral ridges revealed. The latter did not form the lower 
margins of the lateral surface of the nectophore (Fig. 19D). Two 
small patches of cells were found on each side of the nectophore, just 
basal to the vertical lateral ridges, although for many nectophores 
they had been abraded away. In addition there were three distinct 
strips of small epidermal cells, one dorsal and two lateral, stretching 
up from the ostium; the lateral pair being more extensive and 
pronounced than the dorsal one. These were all believed to be sites 
of bioluminescence. Mouth plate absent. Ostiun opened basally. 

The nectosac was Y-shaped in the younger nectophores, but the 
median apical indentation was less pronounced in the larger ones, 





Fig. 20 Parerenna emilyae. Bracts of A. the first and B. the second type. 
Scale 1 mm. 



and may disappear altogether; the nectosac then being T-shaped. 
There was a large muscle-free zone on the apical, adaxial part of its 
lower side. The pallial canal was quite long, extending from the base 
of the thrust block to beyond the point of origin of the pedicular 
canal. The long pedicular canal was inserted onto the nectosac either 
at the point of origin of the lateral radial canals, or slightly basal to 
it. On half of the fully developed nectophores there was a slight 
asymmetry in the origin of the lateral radial canals with either the left 
or the right branching off before the other. The dorsal and ventral 
canals were straight and ran directly to the ostial ring canal. There 
was, however, a slight loop in the lateral radial canals as they curved 
over onto the lateral surface of the nectosac slightly above its mid- 
height. These canals then curved down to the mid-level and continued 
to the ostial ring canal. 

SiPHOSOME. The remaining piece of siphosome was highly con- 
tracted, with four gastrozooids and four palpons still attached. There 
were no signs of any gonodendra. 

Bracts. (Fig. 20). Twenty-two bracts, up to 9 mm in length, 
remain with the specimen. Two basic types, present in approxi- 
mately equal numbers, could be distinguished. The first (Fig. 20A) 
was longer, but narrower, than the second (Fig. 20B). Both types 




Fig. 21 Parerenna emilyae. A. Gastrozooid. Scale 1 mm. B. Tentillum and C. Palpon. Scales 0.5 mm. 



P.R.PUGH 



have two puns of lateral cusps; although in the shorter, more 
rounded bracts, one or both of the more distal pair could be difficult 
to discern. Both types had a patch of small epidermal cells on the 
dorsal surface, but the positioning differed (Fig. 20). Often these 
patches had been abraded away. Proximally the bracteal canal 
originated on the dorsal surface of the bract. It passed down the 
middle o\' the ventral side bract and, for the most part, lay in close 
contact with its surface. A short distance from its distal end it 
narrowed and penetrated through the mesogloea to terminate be- 
neath a hemispherical clump of epidermal cells sunk into the 
mesogloea. Some of these cells, as with the patches on the dorsal 
surface of the bract, probably produced bioluminescence; others 
were nematocysts measuring 80 x 20 |jm. 

Gastrozooid. (Fig. 21 A). Up to 7.5 mm long, with no obvious 
pedicle. The basigaster, to which the tentacle was attached, was 
minute. The gastrodermal lining of stomach region bore a complex 
pattern of villi; while the proboscis was broad and elongate, with a 
distinctive arrangement of eight stripes. 

TENTILLUM. (Fig. 2 IB). The tentillum was very distinctive, with a 
thickened pedicle, occupying about half its length, through which 
the broad gastrovascular canal passed. The distal half of the tentillum 
consisted of an extensive cnidoband and a process, containing a 
canal, that bent away, occasionally at a right-angle, from the base of 
the cnidoband and was terminated by a small spherical swelling. The 
cnidoband appeared to have two rows of large nematocysts on either 
side, measuring 120 x 20 u.m, that, judging by those that had been 
discharged, probably were homotrichous anisorhizas. The remain- 
der of the cnidoband bore numerous smaller nematocysts of two 
sizes; the more cylindrical ones measuring c. 21 x 12 urn, and the 
more ovoid ones c. 26 x 15 um. These were the only nematocysts to 
be found on the distal tip of the cnidoband. 

Palpon. (Fig. 21C). Up to 4 mm long, with a palpacle, without 
nematocysts. at its base. Proboscis region long and narrow with 
broad gastrovascular canal. 

Distribution. Known only from a single specimen collected in 
the region of The Bahamas. 

REMARKS. Although Parerenna emilyae possesses the general 
erennid characters, there are certain differences from those of the 
genus Erenna that warrant its placement in a separate genus. Prima- 
rily, the nectophores are not flattened dorso-ventrally and only the 
apico-lateral ridges are distinct. The weak infra-lateral ridges do not 
demarcate the lower margins of the lateral facets, and the weak 
vertical lateral ridges have an oblique course. Further, the lateral 
radial canals on the nectosac are slightly curved. In addition the 
basigaster of the gastrozooid is minute, especially in comparison 
with the greatly expanded basigasters of the Erenna species. 

ETYMOLOGY. The species is named for my daughter Emily. 



KEY FOR THE IDENTIFICATION OF 
ERENNID NECTOPHORES 

I Nectophores dorso- ventral I v flattened with distinct, short vertical lat- 
eral ridges, and apico- and infra-lateral ridges joining apically. Genus 
Erenna 2 

Nectophores not dorso-ventrally flattened; indistinct vertical lateral 
ridges; indistinct infra- lateral ridges not joining apico-laterals apically. 
Genus Parerenna P. emilyae 



2 Two digitate processes on ventral side of thrust block 
Erenna richardi 



No digitate processes on ventral side of thrust block 3 

3 Apico-lateral ridges divide close to ostium; thrust block with V-shaped 
median indentation; no 'horn' canals Erenna laciniata 

Apico-lateral ridges do not divide close to ostium; thrust block with- 
out median indentation; 'horn' canals usually present 

Erenna cornuta 



Acknowledgements I am extremely grateful to Drs Richard Harbison 
(WHOI) and Edie Widder (HBOI) for inviting me to participate in several 
cruises involving the use of the Johnson-Sea-Link submersibles, and for 
allowing me to use the siphonophore material collected. The reviewers' 
comments were greatly appreciated. I am also grateful to Mike Conquer 
(SOC) for teaching me how to scan the figures. 



REFERENCES 



Alvarino, A. 1969. Zoogeograffa del Mar de Cortes: Quetognatos, Sifonoforos y 
Medusas. Anales del Institute de Biologia, Universidad National Autonoma de 
Mexico. Serie Ciencias del Mar y Limnologia, 40(1). 1 1-54. 

1980. El plancton del Atlantico suroeste. Dinamica y ecologi'a. Boletim do 

lnsti.tu.to Oceanogrdfica, Sao Paulo 29, 15-26. 

1981. Siphonophorae. In 'Atlas del Zooplanclon del Atlantico Sudoccidental' (D. 

Boltovskoy. ed.), pp 383-441. Instituto Nacional de Investigation y Desarrollo 

Pesquero (INIDEP), Argentina. 
Bedot, M. 1904. Siphonophores provenant des campagnes du yacht Princesse-Alice 

(1892-1902). Resultats des Campagnes Scientifiques accomplies par le Prince 

Albert I. Monaco 27, 1-27. 4 pis. 
Bigelow, H.B. 1911. The Siphonophorae. Memoirs of the Museum of Comparative 

Zoology, at Harvard College 38, 173^t02. 
Daniel, R. 1985. Coelenterata: Hydrozoa Siphonophora. The fauna of India and 

adjacent countries. Zoological Survey of India, 440 pp. 
Leloup, E. 1936. Siphonophores calycophorides (suite) et physophorides provenant 

des campagnes du Prince Albert \er de Monaco. Resultats des Campagnes 

Scientifiques accomplies par le Prince Albert I. Monaco 93. 1-36. 

1980. A propos du siphonophore Erenna richardi Bedot. 1904. Bulletin de 

I'InstitUt Royal des Sciences Naturelles de Belgique 52 ( 1 1 ), 1^1. 

Lens, A.D. & van Riemsdijk, T. 1908. The Siphonophora of the Siboga Expedition. 

Siboga-Expeditie (Siboga Expedition) 9, 1-130. 
Margulis, R.Ya. 1969. Distribution of some siphonophore species of the suborder 

Physophorae in the Atlantic Ocean. Vestnik Moskovskogo Universiteta 24, 17-38. 

(In Russian). 
1 977. New data concerning the colony structure in Erenna richardi (Physophorae, 

Agalmidae). Zoologicheskii Zhurnal 56, 148-151. (In Russian). 

1990. Does the species Erenna bedoti (Siphonophora, Physonectae) exist? 

Zoologicheskii Zhurnal 69, 138-142 (in Russian). Translation in Hydrobiological 
Journal 21, 30-34, 1991. 

Moser, F. 1925. Die Siphonophoren der Deutschen Sudpolar-Expedition, 1901-03. 

Deutsche Sudpolar-Expedition 17 (zool 9), 1-541. 
Musayeva, E.I. 1976. Distribution of siphonophores in the eastern part of the Indian 

Ocean. Trudy Instituta Okeanologii 105. 171-197. 
Pugh, P. R. 1975. The distribution of siphonophores in a transect across the North 

Atlantic Ocean at 32°N. Journal of Experimental Marine Biology and Ecology 20, 

77-97. 

1989. Gelatinous Zooplankton - the forgotten fauna. Progress in Underwater 

Science 14, 67-78. 

1998. A re-description of Frillagalma vilyazi Daniel 1966 (Siphonophorae, 

Agalmatidae). Scientia Marina 62. 233-245. 

1999. A review of the genus Bargmannia Totton, 1954 (Siphonophorae, 

Physonecta, Pyrostephidael. Bulletin of the Natural History Museum, London (Zool- 
ogy Series) 65. 51-72. 

Totton, A.K. 1 965. A Synopsis of the Siphonophora. London: British Museum (Natural 
History). 



Bull. not. Hist. Mus. Lond. (Zool.) 67(2): 183-1 



X)C(^2^0G Or \ "\ 



Issued 29 November 2001 



A new species of loach, genus Nemacheilus 
(Osteichthyes, Balitoridae) from Aceh, 
Sumatra, Indonesia 



RENNY KURNIA HADIATY 

Division of Zoology. Research and Development Centre for Biology, The Indonesian Institute of Sciences 
(LIPI). Jin. Raya Jakarta-Bogor, Km. 46, Cibinong, Indonesia 

DARRELL J. SIEBERT 

Department of Zoology, The Natural History Museum, Cromwell Road, London, U.K. SW7 5BD 



SYNOPSIS. A new species of the balitorid genus Nemacheilus is described from Aceh, Sumatra. The new species is 
distinguished from other Nemacheilus species by the combination of a colour pattern of dorsal saddles and lateral blotches but 
without a dark spot at the anterior base of the dorsal fin and the presence a row of enlarged, elongate, posteriorly directed, tear- 
drop shaped scales on either side of the lateral line scale row on the anterior part of the caudal peduncle, each of which bears a 
comparatively large, retrorse, apical tubercle. 



INTRODUCTION 



Little has been reported about fishes inhabiting inland waters of 
Aceh, Sumatra (Kreemer, 1922 (fide Wirjoatmodjo, 1987); Fowler, 
1940; Wirjoatmodjo, 1987; Hadiaty and Siebert, 1998). A small 
collection of fishes was made from Sungai Lembang, Gunung 
Leuser National Park- Aceh Selatan during August-September 1 997, 
as part of an ecosystem mapping project from the Research and 
Development Centre for Biology, Bogor. Gunung Leuser National 
Park, established in 1980, comprises 792,675 hectares of prime 
habitat. It, and an associated management area, include coastal 
lowlands, uplands, and montane habitats. The area is home to tiger, 
elephant, rhinoceros, orangutan and spectacular plants such as 
Rafflesia atjehensis and Amorphophalus sp. The fish collection 
contains several distinctive fishes, of which two species of 
Osteochilus were described earlier (Hadiaty & Siebert, 1998). The 
collection also contains a new species of the baltorid sub-family 
Nemacheilinae that has enlarged and elongate scales bordering the 
lateral line in anterior half of the caudal peduncle, each of which 
bears a large, retrorse tubercle near its posterior tip. 

The Indonesian and Malaysian nemacheiline loaches were last 
revised by Kottelat (1984), who recognised nine species in the region 
but noted that several nominal taxa, especially some from Sumatra, 
could not be critically evaluated because of the lack of sufficient 
material. Since then Chin and Samat (1992) have described N. 
elegantissimus from Sabah, Malaysia and Kottelat etal. (1993) have 
recognised as valid two of the species, TV. pfeifferae and TV. longipinnis, 
that Kottelat could not evaluate earlier. The number of valid species 
recognised species in the region now stands at 12. Two of the species 
in the area, TV. selangoricus and TV. spiniferus, have acuminate scales, 
each of which bears a large tubercle, along the lateral line on the caudal 
peduncle. One, TV. selangoricus, is widely distributed, occurring in 
Sumatra, Malaysia, and Borneo, while the other, TV. spiniferus, is 
known only from Borneo (Kottelat et al., 1993). 



MATERIALS AND METHODS 

Methods for counts and measurements follow Kottelat (1984). The 

© The Natural History Museum, 2001 



specimens of the new species were collected by electrofishing and 
are deposited at the Museum Zoologicum Bogoriense (MZB), Re- 
search and Development Centre for Biology, Cibinong, Java, 
Indonesia and The Natural History Museum (BMNH). London. The 
egg count was done by direct enumeration. Illustrations of scales 
and mouthparts were rendered from camera lucida tracings. Statisti- 
cal testing of differences in the shape of caudal peduncle scales 
between males and females of the new species and between the new 
species and TV. chrysolaimos are based on measurements of camera 
lucida tracing of individual scales made under compound microscopy. 
The abbreviation ZMA is for Zoologisch Museum, Amsterdam. 
Peter Bartsch, Museum fuer Natkurkunde der Humboldt-Uni versitaet 
zu Berlin, examined the holotypes of TV. dunckeri (ZMB 20546) and 
TV. longipinnis (ZMB 20547) for us; he also provided a translation of 
Ahl's (1922) description of the colour pattern of each species. 



SYSTEMATICS 

Nemacheilus tuberigum sp.nov. 

(Figs. 1-5; Tables 1,2) 

Holotype, MZB 9356 (48.5 mm SL, 59.2 mm TL); Indonesia, 
Sumatra, Aceh Selatan, Kecamatan Kluet Selatan, Desa Pucuk 
Lembang, Gunung Leuser National Park, caught in a clear forest 
stream tributary to Sungai Lembang; R.K. Hadiaty and A. Mun'im, 
2 September 1997. 

Allotype, MZB 10565 (43.0 mm SL); same data as holotype. 

Paratypes, MZB 9357, 12 ex., (39.6-53.2 mm SL); same data as 
holotype. BMNH 2000.4.10.1-5, 5 ex., (42.2-50.5 mm SL); same 
data as holotype. MZB 9358, 4 ex., (44.8-53.4 mm SL); same 
location as holotype; R.K. Hadiaty and A. Mun'im, 31 August 1997. 
MZB 9359, 1 ex., (42.6 mm SL); same location as holotype; R.K. 
Hadiaty and A. Mun'im;l September 1997. MZB 9360, 4 ex., 
(42.6-49.2 mm SL) Indonesia, Sumatra, Aceh Selatan, Desa Pucuk 
Lembang, Alur Betung, a tributary of S. Lembang; R.K. Hadiaty and 
A. Mun'im; 2 September 1 997. MZB 936 1 , 2 ex., (3 1 .9 and 37.2 mm 
SL); Indonesia, Sumatra, Aceh Selatan, Suag Balimbing Research 



IS4 



R.K. HAD1ATY AND D.J. SIEBERT 




Fig. 1 A. Holotype of N. tuberigum, MZB 9356. 48.5 mm SL, gravid 
female. B. Allotype of N. tuberigum, MZB 10565, 43.0 mm SL, mature 
male. Scale bar in mm. 

Station, caught in a muddy forest stream tributary to Sungai Lembang; 
R.K. Hadiaty and A. Mun'im; 4 September 1997. 

Non-type materials: MZB 935 1 , 10 ex., (38.0-53.0 mm SL); data 
as for holotype. MZB 9362, 28 ex., (33.2-45.9 mm SL); location as 
for holotype; R.K. Hadiaty and A. Mun'im; 3 1 August 1997. MZB 
9363. 27 ex.. (34.7-52.2 mm SL); location as for holotype; R.K. 
Hadiaty and A. Mun'im; 1 September 1997. MZB 9364, 11 ex., 
(33.2-52.3 mm SL); Indonesia, Sumatra, Aceh Selatan, Desa Pucuk 
Lembang, Alur Betung, a tributary of Sungai Lembang; R.K. Hadiaty 
and A. Mun'im; 2 September 1997. MZB 10566, 2 ex., (47.2-48.2 
mm SL); unnamed tributary of S. Alas, behind the camp at Ketambe 
Research Station, Aceh, Sumatra, Indonesia; R.K. Hadiaty & A. 
Mun'im: 21 June 1998. MZB 10567, 3 ex., (38.6-52.9 mm SL); 
unnamed tributary of S. Alas, Ketambe Research Station, Aceh, 



Sumatra, Indonesia; R.K. Hadiaty & A. Mun'im; 20 June 1998. 
MZB 10568, 5 ex., (41.2-52.8 mm SL); unnamed tributary of S. 
Alas, in front of the National Park camp ground, Ketambe, Aceh, 
Sumatra, Indonesia; R.K. Hadiaty & A. Mun'im; 23 June 1998. 
MZB 10569, 2 ex., (39.8^4.9 mm SL); mouth of S. Sukarimbun, 
Ketambe, Aceh, Sumatra, Indonesia; R.K. Hadiaty & A. Mun'im; 
20 June 1998. MZB 10570, 15 ex., (28.2-59.6 mm SL), unnamed 
tributary of S. Alas, behind the camp of the Ketambe Research 
Station, Sumatra, Indonesia; R.K. Hadiaty & A. Mun'im; 21 June 
1998. 

Diagnosis. Nemacheilus tuberigum is easily distinguished from 
all other Nemacheilus by the following combination of characters: a 
row of comparatively large tubercles present on enlarged, elongate 
scales in the scale rows immediately above and below the lateral line 
scale row on the anterior half of the caudal peduncle, (comparatively 
large tubercles in a similar position present also in N. selangoricus 
and N. spiniferus, but on scales with peculiar posterior elongate 
processes); and colour pattern consisting of 1 1-15 dorsal saddles, a 
series of 8-13 lateral blotches centered along the lateral line and 
without a dark spot at anterior base of dorsal fin (present in N. 
selangoricus and N. spiniferus). 

DESCRIPTION. General appearance and physiognomy are shown 
in Figure 1 ; morphometric and meristic data for the holotype and 
paratypes (range, as the minimum and maximum observation, mean 
and standard deviation) are given in Table 1 . Nemacheilus tuberigum 
presents a shape and colour pattern general for many members of the 
genus. 

Dorsal head length 5-6 times in SL; eye moderate, shorter than 
snout, 3-4 times in dorsal head length; suborbital flaplet present in 
males at anteroventral corner of eye, anterior nares a short tube (Fig. 
2); small tubercles scattered over dorsal and lateral exposures of the 
head. Cephalic lateral line pores: supraorbital canal with 5 pores; post- 
temporal canal with 3 pores, infraorbital canal with 10 pores, supra- 
occipital canal with 3 pores; operculomandibular canal with 9 pores. 

Mouth crescent-shaped (Fig. 3), with three pairs of barbels: 
anterior rostral barbel shortest, reaching to about middle of eye; 
posterior rostral barbel longest, reaching to half way between hind 
edge of eye and hind edge of opercle; mandibular barbel intermediate 




Hy. 2 Lateral view of the head of the allotype of N. tuberigum, MZB 10565. Suborbital flap clearly evident; anterior naris a short tube, valve-like; small 
tubercles are scattered over the head. 



A NEW NEMACHEILUS FROM SUMATRA 



185 




Fig. 3 Mouth of N. tuberigum MZB 9357, male, 48.4 mm SL. Lower lip 
completely separated at the midline; upper lip weakly crenulate, weakly 
papillate in posterolateral part. 

in length between rostral barbels, reaching to end of posterior rostral 
barbel (Fig. 2). Process dentiformis small. Lips moderately fleshy: 
upper lip weakly crenulate to feebly papillose, posterolateral parts of 
upper lip weakly papillose; lower lip with a median incision com- 
pletely separating right and left sides, with 2 or 3 deep pleats on 
either side of median incision. 

Pectoral fin with 11 rays, reaching at most only halfway to 
pelvics; pelvics with 8 rays, nearly reaching anus, anal fin not 
reaching caudal fin. Hindborder of pectoral and pelvic fins with 
prolongations at extremities of rays. Caudal fin forked, short. 

Body fully scaled scales small, ovoid slightly pointed posteriorly. 
Scales on caudal peduncle larger and longer than those from mid- 
body region. Scale rows immediately above and below the lateral 
line on the anterior part of the caudal peduncle with 5-10 obviously 




Fig. 4 Scales of N. tuberigum. A. Scale from mid-body below dorsal fin. 
B. Scale from scale row immediately adjacent to lateral line on caudal 
peduncle. These scales usually bear an enlarged tubercle near the 
posterior margin. Anterior to the left; scale bar = 0.1 mm. 

elongate scales (Fig. 4) (1/w for males x= 1.64, s.d. = 0.055, n = 5; 
1/w for females x = 1.55, s.d. = 0.097, n = 5), usually tear-drop in 
shape, each bearing a large retrorse tubercle (Fig. 5) as on the 
acuminate scales of N. selcmgoricus and N. spiniferus. 




Fig. 5 Close up of anterior caudal peduncle of allotype of N. tuberigum (anterior to the left). Lateral line passes through the upper third of the three oval 
lateral blotches; small tubercles are scattered over the caudal peduncle generally. A row of larger tubercles can be seen on either side of the lateral line, 
beginning near the left edge of the second lateral blotch and ending near the left edge of the third blotch. 



186 



R.K. HAD1ATY AND D.J. SIEBERT 



Lateral line complete, with 76-90 pores, incomplete in specimens 
le-.s than about 30 mm SL. 

In alcohol body pale yellowish with 10-15 irregular dorsal sad- 
dles, saddles w ider than saddle interspaces. There are 8-13 blotches 
along the lateral line, blotches progressively larger posteriorly (Fig. 
lb). In larger specimens blotches may be fused to form longitudinal 
stripe, especially in posterior half of body (9 of 29 specimens with 
such a longitudinal stripe). No spot present at anterior base of dorsal 
fin. Two dark marks at base of caudal-fin rays: the larger, more 
intense mark a vertically elongate dark mark in series with lateral 
blotches; the smaller, a dorsally situated, oval, dark spot, perhaps in 
series with dorsal saddles. 

Sexual dimorphism. In our material, the largest individuals are 
female; no male exceeded 48.5 mm SL while several females 
exceeded this length, the longest of which is 53.4 mm SL. Males 
(39.8mm SL and over) possess a large subocular flaplet (Fig. 2) 
below the anteroventral margin of eye and have a greater number of 
tubercles in the mid-body region than females. In males retrorse 
tubercles are present on each scale in the region between the pelvic 
fin and the lateral line, in well-defined rows. Tubercles are also 
present in females in this region, but they are scattered rather than 
present in rows on every scale; tubercles of females are also smaller 
than those of males. Above the lateral line in the mid-body region 
both sexes have tubercles variously scattered. 



INDIAN 
OCEAN 




o 



>^ 



<? 



98° 



Fi«. 6 Drainage map of Gunung Leuser National Park area from which 
N. tuberigum materials have been captured; the shaded area of the inset 
is the area covered by the drainage map. ▲ = S. Lembang localities near 
Suag Balimbing Research Station; target symbol = type locality. T = S. 
Alas drainage localities near Ketambc Research Station. 



REPRODUCTION. Yellow to orange mature eggs were visible though 
the thin abdominal skin of female with a distended belly. Another 
female, 48.5 mm TL, contained 4192 eggs of uniform size (0.61 mm 
mean diameter). Fishes that spawn all eggs in a single event are 
known as total spawners (Welcomme, 1979). One of the character- 
istics associated with total spawning is uniform egg size. Since eggs 
of A 7 , tuberigum were observed to be of uniform size we interpret the 
species as most likely being a total spawner. 

Distribution and habitat. Nemacheilus tuberigum is known 
only from Sungai Lembang, Suag Balimbing Research Station and 
Sungai Alas, Ketambe Research Station in the region of Gunung 
Leuser National Park, Aceh, Sumatra (Fig. 6). It was taken from 
streams of small to moderate size, mostly in shallow, clear water, but 
a few juveniles were taken from murky water. The substrate over 
which it was found varied from gravely to sandy. Species co- 
occurring with N. tuberigum were: Homaloptera ripleyi, H. 
gymnogaster. Tor tambra, Rasbora lateristriata, R. sumatrana, R. 
meinkenii, Glyptothorax cf. major and G. cf. platypogonoides. 
Given the lack of general information on fish distributions in 
Sumatra it would not be surprising to find the species elsewhere. 

ETYMOLOGY. The epithet tuberigum, a noun, is derived as sug- 
gested in Brown (1956: p. 44). It is constructed in reference to the 
comparatively large tubercles on either side of the lateral line on the 
anterior part of the caudal peduncle; it is from the latin words tuber, 
a neuter noun meaning swelling, and mangus, meaning large, the 
later dismembered to -gus, but modified appropriately to the gender 
of tuber. 



COMPARISONS WITH OTHER SPECIES 



External characters. The presence of a row of enlarged 
tubercles on the caudal peduncle on the apex of elongated scales of 
the scale rows immediately above and below the lateral line scale 
row immediately separates N. tuberigum from all other known 
Nemacheilus except N. selangoricus and N. spiniferus. The lack of 
a dark spot at the anterior base of the dorsal fin immediately 
distinguishes N. tuberigum from these later two taxa; additional 
differences from these two species are recorded in Table 2. Never- 
theless it is worthwhile to consider some general comparisons with 
other Nemacheilus from Sumatra, which include (Kottelat et al., 
1993; Kottelat & Whitten, 1996) N. fasciatus, N. kapuasensis, N. 
lactogeneus, N. longipinnis, and N. pfeiferae and the possibly valid 
nominal taxa N. dunckeri, N. papillosa and N. jaklesii. A third party 
reviewer also requested a comparison to N. masyae, a species 
present in peninsular Thailand and Malaysia at about the same 
latitude as N. tuberigum. 

Colour pattern. The colour pattern of N. tuberigum is a series of 
dorsal saddles and midlateral blotches, a colour pattern general for 
all Sumatra Nemacheilus of which we are aware except N. 
lactogeneus. Nevertheless, N. tuberigum is distinguishable from 
some of the other species of Nemacheilus from Sumatra in matters of 
colour pattern details. 

Nemacheilus fasciatus, N. jaklesii and N. longipinnis each pos- 
sess a dark spot at the anterior base of the dorsal fin, which N. 
tuberigum lacks. In addition, N. jaklesii appears to have a colour 
pattern of dorsal saddles nearly confluent with relatively deep 
vertical, lateral bands rather than blotches along the lateral line. The 
photograph of the type of A/, longipinnis reproduced in Kottelat et al. 
( 1 993; Fig. 1 39) indicates a dark spot at the base of the anterior part 



A NEW NEMACHEILUS FROM SUMATRA 



187 



of the dorsal fin even if the colour pattern is not evident on the rest 
of the body. Peter Bartsch examined this specimen for us and 
confirms the presence of the spot, although now faded, which Ahl 
(1922) specifically mentions in the description of the colour pattern 
of the species. Nemacheilus longipinnis also seems to have a larger 
eye than N. tuberigum, in which the eye is noticeably shorter than 
the snout. 

With 10-15 dorsal saddles and 8- 1 3 lateral blotches, N. tuberigum 
has generally fewer dorsal saddles and lateral blotches than N. 
pfeifferae (10-18 dorsal saddles, 10-15 lateral blotches; data from 
Kottelat et ah, 1993), the photograph of which in Kottelat et al. also 
shows its lateral blotches fused into a more or less continuous band 
from the opercle to the base of the caudal fin. We have examined 2 
syntypes of N. papillosa, the condition of which is not very good. 
However, traces of the colour pattern are still evident; the colour 
pattern seems to consists of dorsal saddles with a well defined, 
relatively narrow, lateral band 3 scale rows wide centered on the 
lateral line, at least on the posterior half of the body. Lateral blotches 
in N. tuberigum may become confluent to form a lateral band, 
especially in the posterior half of the body, but in our material it is 
never as narrow or as distinct as in the syntypes of N. papillosa we 
examined. The colour pattern of some populations of N. kapuasensis 
is readily distinguishable from that of N. tuberigum. Dorsal saddles 
and lateral blotches of these N. kapuasensis are equal in number and 
more or less confluent, rendering a appearance of dorsal saddles that 
extend ventrally to the level of the insertion of the pectoral fin (see 
Kottelat, 1984:Fig. 16b; Kottelat etal, 1993: PI. 25). Dorsal saddles 
and lateral blotches of N. tuberigum form two clearly separate 
series. 

Kottelat et al. (1993) suggest /V. dunckeri may beasynonymof N. 
pfeifferae. The holotype of N. dunckeri was examined for us by 
Peter Bartsch. He reports the specimen to be faded but that there is 
no indication of a dark spot at the anterior base of the dorsal fin; Ahl 
(1922) specifically stated there are no spots on any of the fins. Ahl 
also described a sharply defined dark lateral stripe, wider than eye 
diameter, extending from the opercle to the caudal peduncle. The 
lateral blotches of TV. tuberigum do not form such a band, not even on 
the caudal peduncle of the largest specimens examined by us. 

True N. lactogeneus (contra Kottelat etal., 1993; work in progress) 
are whitish, without any dorsal saddles at all sizes and possess an 
ocellated dark spot at the base of the middle caudal fin rays. Thus, it 
is readily distinguishable from N. tuberigum. 

Nemacheilus tuberigum is easily and immediately distinguish- 
able from N. masyae by colour pattern; N. masyae exhibits a 
prominent dark spot on the anterior, basal part of the dorsal fin 
(Smith, 1933; Kottelat, 1990: Fig. 29) which N. tuberigum does not. 



DISCUSSION 

Squamation and tuberculation among Nemacheilus species varies a 
great deal. The specialised 'acuminate' condition of certain scales, 
with large apical tubercles, on the caudal peduncle of N. selangoricus 
and N. spiniferus as described by Inger & Chin (1962) and Kottelat 
(1984) is perhaps one extreme of a continuum. Nemacheilus 
tuberigum possesses elongate, tear-drop shaped scales, even if not 
attaining a acuminate condition, with comparatively large apical 
tubercles, in the same general area as N. selangoricus and N. 
spiniferus and other species such as N. chrysolaimos and N. papillosa 
also have enlarged or elongated scales on the caudal peduncle, even 
if these species do not possess the distinctive rows of large tubercles 
along the lateral line of the caudal peduncle as do the three species 



just discussed. Of N. tuberigum, N. chrysolaimos and N. papillosa 
the first two have scales on the caudal peduncle that are much more 
elongate than those of N. papillosa and those of N. tuberigum are 
significantly longer than those of N. chrysolaimos (N. tuberigum x = 
1 .59, sd. = 0.08; N chrysolaimos x = 1 .48, sd = 0.06 : t = 3.23, p = 
0.0023). It is conceivable that when more species are examined in 
detail the gaps between different conditions of squamation found 
among species of Nemacheilus will narrow, or even disappear so 
that the transition between conditions will been seen as smooth. 



COMPARATIVE MATERIAL 



Nemacheilus chrysolaimos: MZB 1374 b, 10 ex. (43.0-47.3 mm 
SL); Cisarua, Bogor, Java, Indonesia; Yachya, 5 April 1970. 
MZB 1366, 9 ex. (27.9-41.6 mm SL); Ciapus, Gadog, Bogor, Java, 
Indonesia; Minin, 25 December 1 969. MZB 1 376 b, 3 ex. (45.5-19.4 
mm SL); Cimatuk, Parung Panjang, Bogor, Java, Indonesia; S. 
Wargasasmita, 28 October 1970. 

Nemacheilus fasciatus: MZB 1372 b, 3 ex., (55.3-57.6 mm SL); 
Cikaniki, Cipaku, Bogor, Java, Indonesia; S. Wargasasmita, 25 
March 1970. MZB 1372 c, 6 ex., (56.7-69.2 mm SL); Cikaniki, 
Cipaku, Bogor, Java, Indonesia; S. Wargasasmita, 25 March 1970. 
MZB 2010, 7 ex. (56.3-68.6 mm SL), Sangharus, Airnaningan, 
Pulau Panggung, Lampung Selatan, Sumatra, Indonesia; D. Hardjono 
& F. Sabar, 26 February 1975. 

Nemacheilus jaklesii: BMNH 1866.5.2.60, paralectotype, (49.1 mm 
SL); Paya Kumbuh, Sumatra, Indonesia; male. 

Nemacheilus papillosa: ZMA 112.874, 2 syntypes, (48.0 mm SL, 
male with subocular flap obvious; 56.3 mm SL, female with eggs 
apparent); Lake Toba, Balige, Sumatra, Indonesia. 

Nemacheilus selangoricus: MZB 3551, 3 ex. (28.3-29.4 mm SL); 
small forested stream where it flows into S. Mandai upstream from 
its confluence with Kapuas mainstream, S. Kapuas basin, Kalimantan 
Barat, Indonesia; T.Roberts, 10 August 1976. MZB 2395 b, 3 ex., 
(29.3^-1 .4 mm SL); Tanah merah, Lempake, Kalimantan, Indonesia; 
M. Siluba, 27 February 1978. 

Nemacheilus spiniferus: MZB 6807, 6 ex., (32.2-37.5 mm SL); S. 
Tarusan, a tributary of S. Laung, a tributary of S. Barito, Kalimantan 
Tengah, Indonesia; D.J. Siebert, A.H. Tjakrawidjaja & O. Crimmen, 
16 July 1992. MZB 6877, 11 ex., (29.5-38.0 mm SL); S. Karingian, 
a tributary of S. Laung, a tributary of S. Barito, Kalimantan Tengah; 
D.J. Siebert, A.H. Tjakrawidjaja & O. Crimmen, 7 July 1992. MZB 
6928, 2 ex., (38.7^0.0 mm SL); S. Laung, a tributary of S. Barito, 
Laung Tuhup, Barito Utara, Kalimantan Tengah, Indonesia; D.J. 
Siebert, A.H. Tjakrawidjaja & O. Crimmen, 15-18 July 1992. MZB 
6948, 2 ex., (34.1-34.5 mm SL); S. Mata, a tributary of S. Barito 
below Muara Laung, Laung Tuhup, Barito Utara, Kalimantan Tengah, 
Indonesia; D.J. Siebert, A.H. Tjakrawidjaja & O. Crimmen, 8 July 
1992. 



Acknowledgements. We thank Dr. Soetikno Wirjoatmodjo for reprints 
about a collection he made at Ketambe; Ir. Ike Rachmatika, Drs. Agus H. 
Tjakrawidjaja and Drs. Haryono for their support; and Ir. Daisy Wowor for 
literature. Dr. Peter Bartsch of the Museum fur Naturkunde der Humboldt- 
Universitat zu Berlin kindly examined the holotypes of N. dunckeri and N. 



188 



R.K. HADIATY AND D.J. SIEBERT 



longipinnis for us and provided translations of Ahl's description of (heir 
colour patterns. Dr. Maurice Kottelat provided \ aluable suggestions as a third 
part) re\ iewer. Permission EromTaman Nasional Gunung Leuserand Leuser 
Management Unit to conduct studies in theSuaq Balimhing Research Station 
is grateful]} acknowledged. We also thank the Photography Unit of the 
Natural Histon Museum. London for photographs and Abdul Mun'im for 
dedicated assistance with fieldwork. 



REFERENCES 

Ahl, E. 1422. Einige neue Suesswasserfische des Indo-Malayischen Archipels. 
Sitzungsberichte der GesellschaftNaturforschender Freunde, 1922: 30-36. 

Brown, R. 1956. Composition of Scientific Words, Rev. Ed. 882p. Smithsonian 
Institution Press, Washington, D.C. 

Chin, P.K. & Samat, A. 1992. A new loach. Nemacheilus elegantissimits. (family 
Balitoridae. sub family Nemachelinae) from Danum Valley. Sabah, Malaysia. Ma- 
layan Nature Journal, 46: 25-33. 

Fowler, H.W. 1940. Zoological results of the George Vanderbilt Sumatran Expedition 
1936-1939. Part II -The Fishes. Proceedings of the Academy of Natural Sciences of 
Philadelphia 91: 369-398. 

Hadiaty, R. & Siebert, D. 1998. Two new species of Osteochilus (Teleostei: Cyprini- 



dae) from S. Lembang, Suag Balimbing Research Station. Gunung Leuser National 

Park. Aceh. Northwestern Sumatra. Revue Francaise d'Aquariologie. Herpetologie 

25: 1-2. 
Inger, R. & Chin, P.K. 1962. The Fresh-Water Fishes of North Borneo. Fieldiana 

Zoology 45: 1-268. 
Kottelat, M. 1984. Revision of the Indonesian and Malaysian noemacheiline loaches. 

Japanese Journal of Ichthyology 31: 225-251. 
Kottelat, M. 1990. Indochinese Nemacheilines. A Revision of Nemacheiline Loaches 

(Pisces: Cypriniformes) of Thailand. Burma, Laos, Cambodia and southern Viet 

Nam. 262p. Verlag Dr. Friedrich Pfeii, Munich. 
Kottelat, M., & Whitten, A.J. 1996. Freshwater fishes of Western Indonesia and 

Sulawesi: additions and corrections. 8pp. Periplus Editions, Jakarta. 
Kottelat, M., Whitten, A.J., Kartikasari, S.N. & Wirjoatmodjo, S. 1993. Freshwa- 
ter Fishes of Western Indonesia and Sulawesi, xxxviii + 221 p. 84pl. Periplus 

Editions, Singapore. 
Kreemer, J. 1 922. Atjeh. Algemeen samenvaltend overzicht van land en volk van Atjeh 

en onder hoorigheden. E.J. Brill, Leiden. (Fide Wirjoatmodjo, 1987). 
Smith, H.M. 1933. Contributions to the Ichthyology of Siam. II. New Species of 

Loaches of the Genus Nemacheilus. Journal of the Siam Society, Natural History 

Supplement 9: 53-62. 
Weleomme, R. 1979. Fisheries ecology of floodplain rivers, viii + 317p. Longman, 

New York. 
Wirjoatmodjo, S. 1987. The river ecosystem in the forest area at Ketambe, Gunung 

Leuser National Park, Aceh, Indonesia. Archiv fur Hydrobiologie-Ergebnisse der 

Limnologie 28: 239-246. 



Table 1 Morphometric and meristic data for Nemacheilus tuberigum 
sp.nov., n = 28 for paratypes; measurements follow Kottelat (1984). 





Holotype 


Paratypes 


X 


s.d. 


Standard length 


48.5 


31.9- 


-53.4 






Total length 


122.1 


121.1- 


-127.7 






As % standard length 












Lateral length of head 


22.1 


20.3- 


-24.8 


22.7 


0.96 


Dorsal length of head 


18.6 


16.8- 


-20.9 


19.2 


0.91 


Predorsal length 


48.7 


47.0-52.0 


49 


1.36 


Prepelvic length 


51.2 


47.9- 


-52.8 


50.4 


1.20 


Preanal length 


75.4 


73.0- 


-78.5 


75.7 


1.47 


Preanus length 


69.2 


65.1- 


-73.3 


69.4 


2.11 


Head height (at eye) 


9.6 


9.2- 


11.2 


10.15 


0.42 


Body height (at nape) 


12.2 


11.9- 


-14.5 


12.8 


0.58 


Body height (at dorsal origin) 


17.9 


15.1- 


-22.3 


18.0 


1.73 


Height of caudal peduncle 


10.8 


10.6- 


-11.9 


11.3 


0.33 


Length of caudal peduncle 


14.1 


13.1- 


-16.4 


14.8 


0.74 


Snouth length 


8.8 


7.8- 


-9.8 


8.7 


0.51 


Head width (at nares) 


4.8 


4.5- 


-5.5 


5.0 


0.22 


Maximum head width 


12.9 


12.0- 


-14.5 


7.1 


0.67 


Body width (at dorsal origin) 


12.1 


10.0- 


-16.3 


15.9 


1.55 


Body width (at anal origin) 


6.8 


6.8- 


-8.4 


24.4 


0.38 


Eye diameter 


5.0 


2.3- 


-6.3 


5.0 


0.66 


Interorbital width 


6.3 


6.3- 


-7.9 


7.1 


0.37 


Height of dorsal fin 


15.2 


12.3- 


-20.1 


15.9 


2.03 


Length of upper caudal lobe 


21.9 


21.0-28.5 


24.4 


1.55 


Length of lower caudal lobe 


24.1 


22.0- 


-26.6 


23.6 


1.27 


Length of median caudal ray 


16.8 


16.0- 


-19.8 


17.5 


1.09 


Length of anal fin 


17.4 


15.4-18.2 


16.8 


0.75 


Length of pelvic fin 


15.0 


14.8- 


-17.0 


15.9 


0.65 


Length of pectoral fin 


16.6 


15.7- 


-20.8 


17.8 


1.25 


As % dorsal head length 












Lateral length of head 


118.7 


111- 


-128 


118 


4.52 


Head height at eye 


51.9 


47.5- 


-58.1 


52.9 


2.30 


Head width at nares 


26.0 


23.3- 


-28.1 


26.1 


1.61 


Maximum head width 


69.3 


60.8- 


-76.5 


68.1 


4.88 


Eye diameter 


27.2 


23.9- 


-29.9 


26.5 


1.48 


Interorbital width 


34.1 


32.2^3.1 


36.9 


2.31 


C. peduncle : Length/height 


1.3 


1.1- 


-1.4 


1.3 


0.11 


D. fin raystsimple/branched) 


4/9 


4/9 






Caudal fin rays 


9+8 


9+8 






A. fin rays (simple/branched) 


3/5 


3/5 






Ventral fin rays 


8 


8 






Pectoral fin rays 


11 


II 






Lateral line pores 


82 


76- 


-90 







A NEW NEMACHEILUS FROM SUMATRA 



189 



Table 2 Comparison of selected features helpful in distinguishing Nemacheilus tuberigum from N. selangoricus and N. spiniferus. 



N. tuberigum 



N. selangoricus* 



N. spiniferus" 



Colour Pattern Body with 8-13 dark brown blotches, 

sometimes fused to form a longitudinal 
stripe in larger specimens. 
10-15 dark brown saddles on the back, 
sometimes some of which are in contact 
with blotches on sides. 

Eye diameter Smaller, 5% SL (4.5-5.5%) 

26.5% HL (23.9-29.9%) 

Dorsal fin Dorsal fin dusky, without spot at 

anterior base 
Height of dorsal fin shorter 

15.9% SL (12.3-28.5%) 
85.8% HL (65.8-108.7%) 

Caudal fin Upper caudal lobe, shorter 

24.4% SL (21.0-18.5%) 
126.5% HL (111-146.4%) 
Lower caudal lobe shorter 
23.4% SL (15.7-20.8%) 
123.4% HL (112.8-136.0%) 

Pectoral fin Shorter, 

17.8% SL (15.7-20.8%) 
92.9% HL (83.3-106.2%) 

Lateral line pores Complete, 82.6 (76-90) 



Body with 8-12 dark brown bars. 



Larger, 5.7% SL (4.7-6.9%) 
29% HL (23-33%) 

Dorsal fin with 2 distinct black stripes and 

a black spot at anterior base 

Taller, 

20.2% SL (16.4-24.9%) 

101% HL (81-123%) 

Longer, 

33.4% SL (29.6-40.8%) 

165.0% HL (144 -211%) 

Longer, 

28.8% SL (24.3-33.0%) 

145.0% HL (127-158%) 

Longer, 

21. 1%SL (17.0-24.7%) 

106% HL (92-122%) 

Complete, 84.3 (77-96) 



Body with 10-13 dark brown bars. 



Larger, 6.5% SL (5.7-7.4%) 
31% HL (26-33%) 

Dorsal fin with irregular stripes and a 

black spot at anterior base 

Taller, 

20.7% SL (16.9-23.2%) 

98% HL (80-113%) 

Longer, 

30.6% SL (26.8-34.1%) 

142% HL (120-160%) 

Longer, 

28.8% SL (23.8-34.3%) 

135% HL (112-153%) 

Longer, 

22.3% SL (19.1-26.7%) 

105% HL (94-123%) 

Complete, 88(83-93) 



*Data from Kottelat (1984). 



Bull. not. Hist. Mus. Lond. (Zool.) 67(2):191-207 



X>c/p tf . 0to 4\. \ ^ 



Issued 29 November 2001 



Revision of the western Indian Ocean fish 
subfamily Anisochrominae (Perciformes, 
Pseudochromidae) 



ANTHONY C. GILL 

Fish Research Group, Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 
5BD, U.K. 

RONALD FRICKE 

Ichthyology, Staatliches Museum fur Naturkunde, Schloss Rosenstein, Rosenstein 1, D-70191 Stuttgart, 
Germany. 



CONTENTS 



Introduction 191 

Materials and Methods 192 

Systematics 193 

Anisochrominae Smith 193 

Anisochromis Smith 193 

Anisochromis kertyae Smith 195 

Anisochromis mascarenensis sp.nov 197 

Anisochromis straussi Springer, Smith & Fraser 200 

Key to species Anisochromis 202 

Acknowledgements 202 

References 203 



SYNOPSIS. Monophyly of the Anisochrominae is supported by eight autapomorphies: ectopterygoid and mesopterygoid well 
separated from palatine; preopercle well separated from skull; dorsal insertion of posterior mandibulohyoid ligament; dentary not 
forked; medial origin of A ( section of adductor mandibulae; high number of epineural bones; low number of circumpeduncular 
scales; and low number of lower-limb gill rakers. The subfamily includes a single genus, Anisochromis. with three species, which 
are distinguished on the basis of various meristic and male coloration characters: A. kenyae Smith from east Africa, the Comoros 
Islands and Madagascar; A. mascarenensis sp.nov. from Reunion and Mauritius; and A straussi Springer. Smith and Fraser from 
Saint Brandon's Shoals. 



INTRODUCTION 



The family Anisochromidae was erected by Smith (1954) to accom- 
modate a new genus and species, Anisochromis kenyae, from the 
east coast of Africa. He noted that the species exhibited pronounced 
sexual dichromatism, and proposed a close relationship of the new 
family to the Pseudochromidae. He distinguished the two families 
on the basis of differences in number of vertebrae, head scalation, 
number of lateral lines, gill membrane development, palatine denti- 
tion, fin spine development, fin-ray branching, number of pectoral-fin 
rays, and pelvic-fin position. 

Springer et al. ( 1 977) described a second species of Anisochromis, 
A. straussi, from Saint Brandon's Shoals (= Cargados Carajos) in the 
southwestern Indian Ocean, which they differentiated from A. kenyae 
on the basis of coloration and numbers of dorsal-fin rays, anal-fin 
rays, vertebrae and tubed lateral-line scales. They noted that the 
species is dimorphic, but that eggs were present in both colour 
forms. They therefore examined gonads of the species histologically, 
and concluded it is a protogynous hermaphrodite. They also inves- 
tigated the systematic relationships of the Anisochromidae, and 
proposed that the family is the sister group of the Pseudoplesiopidae, 



and that the two families form a monophyletic group with the 
Pseudochromidae. They therefore synonymised the three families 
under the oldest available name, Pseudochromidae. They noted that 
Anisochromis possessed the following autapomorphies: dorsal- and 
anal-fin spines very weak (versus weak to strong); no scales on head 
(versus head scaled); palatine teeth absent (versus present); fewer 
than four gill rakers on lower limb of first gill arch (versus more than 
seven); branchiostegal membranes with ventroposterior margins 
fused across ventral surface of head (versus separate 
ventroposteriorly); ectopterygoid and mesopterygoid well sepa- 
rated from palatine (versus articulate closely with palatine); and 
ligament from (anterior) ceratohyal attaches to dentary on coronoid 
process (versus at symphysis). 

Godkin & Winterbottom (1985) provided evidence for classifica- 
tion of the Congrogadidae, previously placed in the Blennioidei or 
Trachinoidei, in the Pseudochromidae as the sister group of 
Anisochromis. They relegated the Congrogadidae to subfamilial 
status, along with the Anisochrominae, Pseudochrominae and 
Pseudoplesiopinae. They identified a new autapomorphy for 
Anisochromis (A ] section of the adductor mandibulae originates 
from the preopercle medial to the dorsolateral fibres of the A 2 
section), but noted that several of those reported by Springer et al. 



© The Natural History Museum, 2001 



192 



A.C. GILL AND R. FRICKE 



(1977) were also found in the Congrogadinae, and were thus 
synapomorphies of the two subfamilies: absence of palatine teeth; 
lower gill rakers fewer than seven; bead scaleless or with scales 
confined to the cheek and/or operculum; and gill membranes fused. 
In 1995, the first author and associates made collections of 
shorefishes from Mauritius. Included among the collections was a 
single specimen of an undescribed Anisochromis, which had a 
coloration pattern similar to A. kenyae, but with meristic characters 
more closely resembling A. straussi. A search of museum collec- 
tions re\ ealed a small juvenile specimen of the same species collected 
by J.E. Randall from the adjacent island of Reunion. In 1998, the 
second author made collections of shorefishes from the latter island, 
including nine specimens of the new species. However, he 
misidentified them as A. kenyae and recorded them as such in a 
checklist of the fishes of the Mascarene Islands (Fricke, 1999). 




MATERIALS AND METHODS 



Institutional codes follow Leviton etal. (1985). All measurements to 
the snout tip were made to the midanterior tip of the upper lip. 
Length of specimens are given in mm standard length (SL), which 
was measured from the snout tip to the middle of the caudal 
peduncle at the vertical through the posterior edge of the dorsal 
hypural plate. Head length was measured from the snout tip to the 
posteriormost edge of the opercular membrane. Snout length was 
measured over the shortest distance from the snout tip to the orbital 
rim, without constricting the fleshy rim of the latter. Orbit diameter 
was measured as its fleshy horizontal length. Interorbital width was 
measured as the least fleshy width. Upper jaw length was measured 
from the snout tip to the posterior edge of the maxilla. Predorsal, 
preanal and prepelvic lengths were measured from the snout tip to 
the base of the first spine of the relevant fin. Body width was 
measured between the posttemporal pores (Fig. 1). Caudal peduncle 
length was measured from the base of the last anal-fin ray to the 
ventral edge of the caudal fin at the vertical though the posterior 
edge of the ventral hypural plate. Caudal peduncle depth was 
measured obliquely between the bases of the last dorsal- and last 
anal-fin rays. Measurements of fin rays excluded any filamentous 
membranes. Pectoral fin length was measured as the length of the 
longest middle ray. Caudal fin length was measured as the length of 
the lowermost ray on the dorsal hypural plate. 

The last ray in the dorsal and anal fins is divided at its base and 
was counted as a single ray. As in most actinopterygians, the 
uppermost ray in the pectoral fins is rudimentary and rotated so that 
the asymmetrical medial and lateral hemitrichia appear to represent 
two separate rays; these were counted as a single ray. Procurrent 
caudal-fin ray counts were of the rays above ('upper') and below 
('lower') the principal caudal-fin rays. The uppermost principal 
caudal-fin ray was defined as the ray articulating with hypural 5, and 
the lowermost principal caudal-fin ray was the ray articulating with 
the cartilage nubbin between the distal tips of the parhypural and the 
haemal spine of preural centrum 2 (= post-haemal spine cartilage of 
PU2 following the terminology of Fujita, 1989). Counts of tubed 
scales in the anterior lateral lines included both intermittent non- 
tubed scales and empty scale pockets; if the scale following the last 
tubed scale in the anterior lateral line was missing, a value for the 
anterior lateral line count was not recorded. 'Scales in lateral series' 
was defined as the number of scales in the anterior lateral line plus 
the number of scales rows on the caudal peduncle, the latter count 
beginning with the transverse row following the last tubed scale in 
the anterior lateral line and finishing with the transverse row passing 



B 



ATP 



ALLS1 



PARP 



POTP 



SOTP 



NASP 




DENP 




DENP 



POPP 



Fig. 1 Cephalic laterosensory pores of Anisochromis mascarenensis, 
SMNS 23037, 23.9 mm SL, male, holotype, Reunion, in A) dorsal, B) 
left lateral and C) ventral views. AIOP, anterior interorbital pores; 
ALLS I , first anterior lateral-line scale (shown only in B; other scales 
omitted); AN. anterior nostril; ATP, anterior temporal pore; DENP, 
dentary pores; ITP, intertemporal pores; NASP, nasal pores; PARP, 
parietal pores: PIOP, posterior interorbital pores; PN, posterior nostril; 
POPP, preopercle pores; POTP, posterior otic pores; PTP, posttemporal 
pore; SOBP. suborbital pores; SOTP, suprotic pores. Arrow indicates 
dorsal tip of preopercle; branchiostegal membranes shown hatched; 
scaled areas on body shown in manual stipple. Scale bar = 1 mm. 



REVISION OF ANISOCHROMINAE 



193 



through the scale at the midposterior edge of the hypural plate. 
'Scales in transverse series' were counted anterodorsally from the 
anal-fin origin to the dorsal-fin base and are presented in the form 'x 
+ 1 + y = z' where 'x' is the number of scales between the anterior 
lateral line and the anal-fin origin, T is the anterior lateral-line 
scale, 'y ' is the number of scales above the anterior lateral line to the 
dorsal-fin base, and 'z' is the total number of scales in the series. 
Scale-row number for the position of the first ctenoid scale was 
determined by aligning the scale along an anterodorsal scale row to 
the anterior lateral line, then recording the number (counting from 
the origin of the lateral line) of the tubed scale in that row. Gill-raker 
counts were of the outer rakers on the first arch; the angle raker is 
included in the lower-limb (second) count. Counts of pseudobranch 
filaments included all rudiments. Because counts of gill rakers and 
pseudobranch filaments could not be made without damaging the 
branchiostegal membranes, these were only made on a few speci- 
mens. 

Nomenclature of cephalic laterosensory pores is clarified in Fig. 
1 , and follows Winterbottom ( 1 986), except for the nomenclature of 
those of the posttemporal area, which follows Gill et al. (2000). 

Vertebral counts are given in the form precaudal + caudal = 
total. Caudal vertebrae are defined as those with a haemal spine, 
and include the terminal urostylar complex (which was counted as 
a single vertebra). The pattern of insertion of supraneural 
(predorsal) bones and anterior dorsal-fin pterygiophores within 
interneural spaces is given as an 'anterior dorsal-fin pterygiophore 
formula' modified from the 'predorsal formula' of Ahlstrom et al. 
(1976). Each supraneural is represented by an 'S,' neural spines 
are represented by slashes, and pterygiophores are represented by 
'2' (indicating a pterygiophore that bears a supernumerary ray 
and a serially associated ray), or '1' (indicating a pterygiophore 
that bears only a serially associated ray). A superscript V indi- 
cates where a supraneural bone was present only as a vestige. 
Parentheses enclose elements that may be absent. An 'anterior 
anal-fin pterygiophore formula' is also presented, where the 
slashes represent haemal spines. Epineural counts are of the inter- 
muscular bones that have been traditionally identified as epipleural 
ribs by perciform workers; this terminology follows Johnson & 
Patterson (1993: 557, 'the bones conventionally called epipleurals 
in those fishes [afherinomorphs and perciforms and their rela- 
tives] are homologous with the epineurals, not the epipleurals, of 
non-acanthomorph fishes'). Posterior epineurals were often poorly 
ossified, and difficult to count accurately on radiographs; counts 
should therefore be considered approximate. Osteological features 
were determined from radiographs and from cleared-and-stained 
specimens, which were prepared following the methods of Taylor 
& Van Dyke (1985). Examination of ligaments was enhanced by 
transfer of cleared-and-stained specimens from glycerol to 70% 
ethanol. 

We use the terms 'female' and 'male' in preference to 'ocellated 
phase' and 'terminal phase' of Springer etal. (1977). We appreciate, 
however, the deficiencies of this terminology. As noted by Springer 
et al.(p. 7) for A. straussi: 'specimens with ocellated stage colour 
pattern are smaller and generally females, and specimens with 
terminal stage colour pattern are larger and are males, but there is 
considerable overlap. The overlap is not surprising and is compar- 
able to that found in other transforming hermaphrodites.' 

Count and morphometric value ranges are given first for all 
specimens, followed, where variation was noted, by values for the 
holotype in parentheses; where bilateral counts were recorded from 
the holotype, both counts are given and separated from each other by 
a slash, the first count being the left count. Frequency distributions 
for selected meristic characters are summarised in Tables 1-11. 



SYSTEMATICS 



ANISOCHROMINAE SMITH, 1954 
Anisochromidae Smith, 1954: 298. 

Anisochromis Smith, 1954 

Anisochromis Smith 1954: 298 (type species, Anisochromis kenyae 
Smith, 1954, by original designation and monotypy). 

MONOPHYLY 

The following eight autapomorphies diagnose Anisochromis as 

monophyletic: 

1. Ectopterygoid and mesopterygoid well separated from pcdatine. 
In pseudochromines and pseudoplesiopines the palatine is loosely 
connected to the ectopterygoid via a short ligament and to the 
mesopterygoid via a short, narrow ribbon of cartilage. In 
congrogadines the palatine is closely applied, and often tightly 
bound, to the mesopterygoid; the ectopterygoid lacks an anterior 
process, and is disassociated from the dorsal part of the 
palatoquadrate (e.g., Godkin & Winterbottom, 1985: fig. 6; Gill 
et al., 2000: fig. 3). In anisochromines the palatine is broadly 
separated from the ectopterygoid and mesopterygoid: the 
ectopterygoid is truncated anteriorly (though closely applied to 
the mesopterygoid), and lacks a ligamentous connection to the 
palatine; the mesopterygoid is connected to the palatine via a 
relatively long, broad, strap-like ligament (which lacks cartilage) 
(Fig. 2). 

2. Preopercle well separated from skull. In congrogadines, pseudo- 
chromines and pseudoplesiopines the dorsal tip of preopercle is 
in close proximity to the skull, and the preopercular laterosensory 
canal communicates with the pterotic canal via a short membra- 
nous canal; the dorsal tip of the preopercle reaches to near the 
dorsal margin of the hyomandibula (e.g., Gill etal., 2000: figs 1, 
3). In anisochromines the preopercle is truncated dorsally, not 
reaching the dorsal margin of the hyomandibula (reaching to or 
slightly above the opercular process of the hyomandibula), and 
well-separated from the skull (Fig. 2); the preopercular 
laterosensory canal communicates with the pterotic canal via a 
relatively long membranous canal. 

3. Dorsal insertion of posterior mandibulohyoid ligament. Springer 
et al. (1977) proposed that Anisochromis is autapomorphic in 
having a modified orientation of a cord-like ligament extending 
from the anterior ceratohyal to the dentary, from insertion near 
the symphysis (pseudochromines and pseudoplesiopines) to in- 
sertion on the coronoid process. However, we present a different 
interpretation of this character. There are actually two more-or- 
less cord-like ligaments connecting the dentary and anterior 
ceratohyal in anisochromines, pseudochromines and 
pseudoplesiopines. [We use the general term 'mandibulohyoid' 
for these ligaments following Greenwood (1995), though we do 
not intend to imply homology with the mandibulohyoid liga- 
ments of lower teleosts.] The first of these extends from the 
medial or dorsal surface of the anterior ceratohyal, in the vicinity 
of a dorsal notch in the bone (possibly homologous with the 
beryciform foramen of McAllister, 1968), to the dentary sym- 
physis. The second mandibulohyoid ligament extends from the 
lateral surface of the anterior ceratohyal at a point just anterior to 
the anterior/posterior ceratohyal suture to either the posterior end 
of the ventral process (pseudochromines and pseudoplesiopines) 



194 



A.C. GILL AND R. FRICKE 



HYOM p 




DEN 



INTHY 



ANGART 



RETART 



IOP x pop 



QUAD 



Fig. 2 Lateral view of lower jaw and suspensorium of Anisochromis kenyae, RUSI 4906. 23.3 mm SL, right side reversed. ANGART - angularticular; 
DEN - dentary: ECTPT - ectopterygoid; HYOM - hyomandibula; INTHY, interhyal; IOR interopercle; LIG - ligament between palatine and 
mesopterygoid; MESPT. mesopterygoid; METPT, metapterygoid; OP, opercle; PAL, palatine; POP. preopercle; QUAD, quadrate; RETART, retro- 
articular; SOP. subopercle; SYM, symplectic. Cartilage shown in coarse stipple. Scale bar = 1 mm. 



CHL 



PCH 




SMHL 



DHH' 



ACH' 



Fig. 3 Medial view of right side dentary and hyoid bar with associated 
ligaments of Anisochromis kenyae, RUSI 4906, 23.3 mm SL. ACH, 
anterior ceratohyal; BR, branchiostegal rays; CHL, ligament extending 
between posterior and anterior ceratohyals; DEN, dentary; DHH, dorsal 
hypohyal; PCH, posterior ceratohyal; PMHL. posterior mandibulohyoid 
ligament (portion obscured by anterior ceratohyal shown in broken 
lines); SMHL. symphyseal mandibulohyoid ligament; VHH. ventral 
hypohyal. Cartilage shown in coarse stipple. Scale bar = 1 mm. 

or the coronoid process (anisochromines) of the dentary (Fig. 3). 
We interpret this change in orientation as an autapomorphy of the 
Anisochrominae. Although the former (symphyseal) ligament is 
well-developed in congrogadines examined by us (including 
species of Blennodesmus Giinther, Congrogadus Giinther, 
Halidesmus Giinther and Haliophis Riippell ) and in the basalmost 
genus Rusichthys Winterbottom (R.D. Mooi, pers. comm.), the 
latter (posterior) ligament is apparently absent (which may thus 
represent a synapomorphy of the Congrogadinae). 
4. Dentary not forked. In congrogadines, pseudochromines and 
pseudoplesiopines, the dentary is distinctly forked posteriorly, 
with a relatively narrow lamina of bone connecting the promi- 
nent coronoid and ventral processes; this condition is typical of 
perciform fishes. In anisochromines, the coronoid and ventral 
processes arc united by a broad lamina of bone, so that the 
posterior margin of the dentary is weakly concave to almost 
vertical (Figs 2. 3). 



Medial origin of Al section of adductor mandibulae. In pseudo- 
chromines, pseudoplesiopines and most congrogadines, a 
strap-like A : section of the adductor mandibulae originates from 
the vertical limb of the preopercle, overlying other cheek muscu- 
lature (Godkin & Winterbottom, 1985: figs 1A, IB, 1C, 2A and 
2B). In some derived congrogadines, the A section is fused to the 
A, section (Godkin & Winterbottom, 1985: fig. 2C). Aniso- 
chromines have a strap-like A t section, but it originates on the 
vertical limb of the preopercle medial to the dorsolateral fibres of 
the A 2 section (Godkin & Winterbottom, 1985: fig. ID). 
High number ofepineural bones. Epineural bones are present on 
all precaudal vertebrae, and, depending on species, on caudal 
vertebrae 1 through 2-8 in pseudochromines and pseudoplesio- 
pines (Gill, in press; Gill & Edwards, in press). Taken in the 
context of Gill's (1998) interpretation of intermuscular homo- 
logy in congrogadines, Godkin & Winterbottom's (1985) 
descriptions indicate that, depending on species and specimen 
size, epineural bones may be confined to the first few precaudal 
vertebrae, or 'present as far back as the first few caudal vertebrae' 
in congrogadines. In anisochromines, epineural bones are present 
on all precaudal vertebrae, extending posteriorly on to the first 
10-17 caudal vertebrae. 

Low number of circumpeduncular scales. Depending on species, 
pseudochromines and pseudoplesiopines usually have 16 or 
more circumpeduncular scales; although two pseudochromine 
species (Cypho zaps Gill and Pseudochromis striatus Gill, Shao 
and Chen) may have as few as 14 circumpeduncular scales, the 
modal count for both is 16 (Gill, in press). Circumpeduncular 
counts are not obtainable in congrogadines, as the dorsal and anal 
fins are confluent with the caudal fin (fin condition not deter- 
mined for the basal congrogadine Rusichthys plesiomorphus 
Winterbottom, but confluent in its congener R. explicitus 
Winterbottom; Winterbottom, 1 996); nevertheless, congrogadines 
have numerous, small scales on the caudal peduncle. In 
anisochromines, the scales on the caudal peduncle are relatively 
large, with only 12-14, modally 12, circumpeduncular scales. 
Low number of lower-limb gill rakers. Numbers of outer rakers 
on the lower limb of the gill arch (those on ceratobranchial 1 - 



REVISION OF ANISOCHROMINAE 



195 



including the so-called angle raker - and hypobranchial 1 ) range 
from 9-17 in pseudochromines, 6-20 in pseudoplesiopines, and 
5-15 in congrogadines. Anisochromines only have 2-3 lower- 
limb rakers, although 1-2 very tiny rudiments may be present 
ventral to these; all rakers and rudiments are restricted to the 
upper part of ceratobranchial 1 . A low number of lower-limb 
rakers was initially proposed as a character of Anisochrominae 
(as Anisochromidae) by Smith (1954), and cladistically inter- 
preted as an autapomorphy of the taxon by Springer et al. (1977). 
It was later rejected as an autapomorphy of Anisochrominae by 
Godkin & Winterbottom (1985) and interpreted instead as a 
synapomorphy of the Anisochrominae + Congrogadinae. This 
interpretation is problematic because counts for congrogadines 
broadly overlap those of pseudoplesiopines. In any case, lower- 
limb gill-raker counts for anisochromines are lower than has 
been observed in any congrogadines (some descriptions give 
counts as low as 4 for certain congrogadine species, but these 
exclude the angle raker), and we therefore interpret the very low 
number of lower-limb rakers in anisochromines as autapomorphic. 

Additional diagnostic characters 

Additional characters useful in distinguishing anisochromines from 
other pseudochromids are the following: dorsal-fin rays 1,25-27, all 
or all but first segmented rays branched; anal-fin rays 1,17-19, all 
segmented rays branched; pectoral-fin rays 13-15; pelvic-fins rays 
1,4, medial ray small, inconspicuous and unbranched, all other 
segmented rays branched; caudal-fin rays 4-6 + 8 + 8 + 3-5 = 23- 
27; vertebrae 10 (rarely 11) + 22-25; head without scales (predorsal 
scales extending anteriorly to point ranging from about 2/3 distance 
from dorsal origin to parietal commissure, to just short of parietal 
commissure; Fig. 1 ); lateral line represented on body by anterodorsal 
series of 28-39 tubed scales, and posterolateral series of centrally 
pitted scales; parietal pores relatively numerous (total pores 13^44), 
in continuous or almost continuous series over top of head; lower lip 
complete (uninterrupted at symphysis) with deep symphyseal notch; 
branchiostegal (gill) membranes broadly united, but free from isth- 
mus (Fig. 1); fin spines weak and flexible; and anterior dorsal-fin 
pterygiophore formula S/S/(S V or S) + 2/1 + 1. 

Remarks 

We here consider the gender of Anisochromis to be feminine, in 
keeping with the accepted gender of Chromis [see Opinion 1417 
(International Commission on Zoological Nomenclature, 1986) for 
ruling on the gender of Chromis]. Without explanation, Eschmeyer 
& Baily (1990: 29) gave the gender of Anisochromis as masculine; 
presumably their conclusion of masculine gender relates to the 
proposal by Bailey et al. ( 1 980) to have all generic names ending in 
-chromis to be ruled as masculine. 

Etymology 

The generic name is a combination of the Greek anisos, meaning 
'unequal' or 'different,' and Chromis, a genus of pomacentrid fish. 
Gender is feminine (see Remarks above). 

Anisochromis kenyae Smith, 1954 

African Annie 

Figs 2-7, 8A; Tables 1-11 

Anisochromis kenyae Smith, 1954: 300, fig. 1, pi. 6 [Type locality: 
Malindi, Kenya; holotype RUSI 149].-Smith, 1977: 22, pi. 4C, D 
[taxonomic notes; range extension; colour illustrations]. -Springer 
etal., 1977: 5, fig. lc [comparison]. -Wheeler, 1985: 113 [compi- 
lation; text fig.].-Smith, 1986: 539, pi. 46, fig. 169.1A-B 
[compilation; colour illustrations].-Gill, 1998: fig. 5 [osteologi- 
cal details] -Gill & Edwards, 1999: fig. 8A [osteological details]. 



Diagnosis 

The following characters distinguish A. kenyae from congeners: 

dorsal-fin rays 1,25-26, usually 1,25; anal-fin rays 1,17-18, usually 

1,17; caudal vertebrae 2224, usually 23; scales in lateral series 37- 

44, usually 38^41 ; and anterior lateral-line scales 28-35, usually 

30-34. 

Description (based on 46 specimens, 13.8-25.6 mm SL) 
Dorsal-fin rays 1,25-26 (1,25), all or all but first segmented rays 
branched (all branched in holotype); anal-fin rays 1,17-18 (1,17), all 
segmented rays branched; pectoral-fin rays 13-15 (14/14), upper 1- 
2(1/1) and lower 0-1 (1/1) rays simple; pelvic-fin rays 1,4, medial 
ray small, inconspicuous and unbranched, all other segmented rays 
branched; upper procurrent caudal-fin rays 4-6 (5); lower procurrent 
caudal-fin rays 4-5 (4); principal caudal-fin rays 8 + 8, upper 0-1 (0) 
and lower 0-2 (0) unbranched; total caudal-fin rays 24-27 (25); 
scales in lateral series 31-AA (38/38); anterior lateral-line scales 28- 
35 (31/32); anterior lateral line terminating beneath segmented 
dorsal-fin ray 17-21 (20/21); predorsal scales 4-8 (6); scales in 
transverse series 10-13 + 1 + 2-3 = 14-17 (10 + 1 + 2/10 +1+2); 
circumpeduncular scales 12-13; gill rakers 1-2 + 2-3 = 3-5, some- 
times with 1-2 tiny rudiments (inconspicuous ossifications bearing 
a few or no teeth) above and below rakers (gill raker count not 
determined in holotype); pseudobranch filaments 6-7 (not deter- 
mined in holotype). 

Cephalic laterosensory pores (all bilaterally paired, unless 
otherwise stated): nasal pores 2-3 (2/2); anterior interorbital pores 
2-3 (2/2); posterior interorbital pores (unpaired) 1-5 (2); supraotic 
pores 2-6 (5/3); suborbital pores 9-14 (10/10); posterior otic pores 
1-7 (5/5); preopercular pores 8-17 (17/15); dentary pores 3^1 (4/4); 
intertemporal pores 1-2 (1/1); anterior temporal pores 0-1 (1/1); 
posttemporal pores 1-2 (2/1); total parietal pores 13 — 43 (36). 

As percentage of standard length (based on 21 specimens, 18.5— 
24.5 mm SL): body depth at dorsal-fin origin 19.6-23.0 (22.5); 
greatest body depth 20.8-24.4 (23.0); head width 11.8-14.1 (14.1); 
head length 25.5-28.5 (28.2); snout length 4.6-5.6 (4.7); orbit 
diameter 7.1-8.7 (8.5); interorbital width 3.2^4.9 (4.2); upper jaw 
length 8.4-10.3 (10.3); caudal peduncle depth 13.0-15.5 (15.5); 
caudal peduncle length 8.0-10.6 (8.0); predorsal length 29.5-32.9 
(32.9); preanal length 49.7-54.1 (54.0); prepelvic length 22.1-26.3 
(26.3); first segmented dorsal-fin ray length 5.5-10.9 (9.4); third- 
from-last segmented dorsal-fin ray length 1 2.2-15.9 ( 14. 1 ); dorsal-fin 
base length 60.5-66.5 (62.0); first segmented anal-fin ray length 
5.6-10.0 (5.6); third-from-last segmented anal-fin ray length 12.9- 
14.8 (13.6); anal-fin base length 34.3^10.4 (37.1 ); caudal-fin length 
19.6-22.6 (21.6); pectoral-fin length 17.3-21.7 (18.8); pelvic-fin 
length 12.9-17.4(15.0). 

Lower lip complete with deep symphyseal notch; fin spines weak 
and flexible; anterior dorsal-fin pterygiophore formula S/S/(S V or S) 
+ 2/1 + 1 (S/S/2/1 + 1); 20-23 (22) consecutive dorsal-fin 
pterygiophores inserting in 1 : 1 relationship directly behind neural 
spine 4; anterior anal-fin pterygiophore formula 2/1 + 1 or 12 + 1 + 
1 (2/1 + 1); 12-15 (14) consecutive anal-fin pterygiophores insert- 
ing in 1:1 relationship directly behind haemal spine 2; fourth 
segmented pelvic-fin ray longest; caudal fin rounded; ctenoid scales 
beginning at 1-7 (not determined for holotype) transverse scale 
rows behind branchial opening; dorsal and anal fins without distinct 
scale sheaths, though often with intermittent scales overlapping fin 
bases; intermittent series of centrally pitted scales originating on 
midside above anterior part of anal fin, extending posteriorly along 
caudal peduncle to middle part of caudal-fin base; additional 1-3 
centrally pitted scales present above and below pitted scale(s) on 
middle part of caudal-fin base; cheeks, operculum and upper part of 



ls>6 



A.C. GILL AND R. FRICKE 



head without scales: predorsal scales extending anteriorly to point 
ranging from about 2/3 distance from dorsal origin to parietal 
commissure, to just short ol parietal commissure: vertebrae 10 + 22- 
24 ( 10 + 22): epuials 2: epineurals present on vertebrae 1 through 
20-23 (21); pleural ribs present on vertebrae 3 through 10. the 
ultimate rib small to moderately developed. 

Upper jaw with 3 or 4 (at symphysis) to I or 2 (on sides of jaw) 
irregular rows of small conical teeth, those of outer row much larger; 
lower jaw with 2 or 3 (at symphysis) to 1 (on sides of jaw) rows of 
small conical teeth, those of outer row much larger; vomer with I or 
2 rows of small conical teeth arranged in chevron; palatines eden- 
tate; tongue edentate and moderately pointed. 

Live coloration 

Males (based on the description in Smith, 1954: 302, the colour 
illustration in Smith. 1977: pi. 4D, and a photograph of a specimen 
from the Comoros Islands, ROM 56501. 22. 1 mm SL): head bright 
reddish orange to bright red, with black-edged white stripe extend- 
ing from posterodorsal rim of orbit, above upper part of preopercle, 
to upper edge of operculum; white spot on posteroventral rim of 
orbit at about 3 o'clock position; narrow brown to dark grey bar 
extending from ventral part of orbital rim to posterior edge of 
maxilla; iris yellow, red centrally, with radiating brown bars; nape 
dark brownish red to dark grey, with scattered small white spots; 
body black, with scattered small white spots, these sometimes 
aligning to form vague bars on upper part of body; dorsal fin with 
large dark grey to black spot, extending from first segmented ray to 
about fourth or fifth segmented ray; black spot bordered anteriorly 
with yellow, basally with bright yellow to bright orange, and some- 
times posteriorly with yellow to bright orange; basal one-third of 
dorsal fin behind large black spot bright red, with small black spot or 
streak at base of each of fin ray, these sometimes edged anteriorly 
with white; remainder of dorsal fin reddish or pinkish hyaline to 
bright red, with greyish hyaline to grey distal margin; basal one- 
third of anal fin bright red, with small black spot or streak at base of 
each of fin ray, these sometimes edged anteriorly with white; 
remainder of anal fin reddish or pinkish hyaline to bright red, with 
greyish hyaline to grey distal margin; caudal fin black basally, 
remainder of fin greyish hyaline to black; pectoral fin black with 
irregular white spots basally, remainder of fin greyish hyaline, with 
fin rays dark grey; pelvic fin bright yellow on base, remainder of fin 
black, sometimes with distal margin pale grey to hyaline. Females 
(based on the description in Smith, 1954: 302, the colour illustration 
in Smith, 1977: pi. 4C, and a photograph of a specimen from the 
Comoros Islands, ROM 56502, 22.7 mm SL): head olive-brown to 
brown dorsally, becoming pale green ventrally, with two pale olive 
bars on nape; large dark grey to black spot on subopercle, bordered 
irregularly with white, sometimes with additional, smaller white- 
edged black spot on lower part of subopercle; brown-edged diffuse 
pale olive stripe extending from posterodorsal rim of orbit, above 
upper part of preopercle, to upper edge of operculum; two white 
spots or clusters of white to mauve spots on posteroventral rim of 
orbit, at about 3 and 5 o'clock positions; head and nape with 
scattered white to mauve or pale olive small spots; narrow dark 
brown to grey bar extending from ventral part of orbital rim to 
posterior edge of maxilla; iris pale yellow to pale orange, with 
radiating brown bars; body generally orange-brown, becoming ol- 
ive to dusky green posteriorly; body with olive-brown bars, 
alternating with pale olive bars; pale olive bars sometimes dotted 
with pale green to mauve spots; broad orange-yellow area some- 
times present on side of body: upper and lower edges of caudal 
peduncle sometimes with small, punctate black spots; dorsal and 
anal fins dusky green to dusky orange or greenish hyaline, often with 




Fig. 4 Anisochromis kenyae, ROM 567 11, 21.6 mm SL, male, Comoros 
Islands (Photograph by R Hurst). 

irregular pale olive to mauve or white small spots; barring on body 
sometimes extending slightly on to fin bases; base of each ray in 
dorsal and anal fins sometimes with small reddish brown to black 
spot or streak, edged anteriorly with pale olive to mauve or white; 
caudal fin olive to lime green basally, remainder of fin dusky olive 
to dusky orange or hyaline; fleshy pectoral-fin base maroon to dark 
brown, with scattered white to mauve spots; pectoral fin orangish 
hyaline to hyaline; pelvic fin olive, sometimes with base pale 
yellow. 

Preserved coloration 

Males (Figs 4, 8A): pattern generally similar to live coloration, head 
and anterior part of body becoming pale brown, paler ventrally; pale 
markings on head obsolete; dark spots and stripes on head remain, 
becoming dark grey-brown to dark brown; body behind pectoral-fin 
base dark grey-brown; white spots and bars on body remain, becom- 
ing brownish white to pale yellow; dorsal and anal fins brownish 
white, sometimes dusky hyaline distally, with dark grey-brown 
spots often present at base of each fin ray; large dark spot at anterior 
of dorsal fin remains, becoming dark grey-brown; caudal fin dark 
grey-brown basally, dusky hyaline to brown distally; pectoral fin 
grey-brown basally, dusky hyaline to hyaline on remainder of fin; 
pelvic fin brownish white to pale yellow basally, sometimes slightly 
darker on extreme base, remainder of fin dark grey-brown to brown, 
often with distal margin narrowly pale brown. Females (Fig. 5): 
pattern generally similar to live coloration, head and body becoming 
pale brown; pale markings on head and body obsolete; dark spots 
and stripes on head remain, becoming dark grey-brown to dark 
brown (ocellated spots sometimes absent, although possibly these 
represent intermediately coloured, sex-transforming specimens), 
sometimes with additional irregular brown vermiform markings and 
spots on cheek and operculum; dark barring on head and body 
variably remains, becoming brown to pale brown; dark punctate 
spots on caudal peduncle remain, becoming dark brown to dark 
grey-brown, sometimes extending anteriorly to near middle of 
dorsal- and anal-fin bases; dorsal and anal fins dusky brown to 




Fig. 5 Anisochromis kenyae, ROM 56502, 22.7 mm SL, female. 
Comoros Islands (Photograph by P. Hurst). 



REVISION OF ANISOCHROMINAE 



197 



brownish hyaline, often with irregular pale brown small spots; 
barring on body sometimes extending slightly on to fin bases; caudal 
fin brown to grey brown, remainder of fin dusky brown to brownish 
hyaline; fleshy pectoral-fin base brown, with indistinct scattered 
pale spots; pectoral fin brownish hyaline to hyaline; pelvic fin dusky 
brown to brownish hyaline, sometimes paler ventrally, usually pale 
brown to hyaline distally. 

Habitat and Distribution 

Anisochromis kenyae is known only from the east coast of Africa 
(Kenya to northern Mozambique), the Comoros Islands and north- 
ern Madagascar (Fig. 6). As noted above, Fricke's ( 1999) record of 
the species from Reunion is based on specimens of A. mascarenensis. 
According to Smith ( 1954: 302), type specimens of A. kenyae were 
collected from coastal 'pools in reefs at about low-tide mark' and 
that they were 'not uncommon at some localities.' Data accompany- 
ing subsequently collected specimens indicate that the species also 
occurs on reef crests and in shallow subtidal reef areas to depths of 
at least 3 m. 

Comparisons with other species 

The three species of Anisochromis differ from each other in the 
following meristic characters: segmented dorsal-fin rays (modally 
25 in A. kenyae versus modally 26 in A. mascarenensis and A. 
straussi; Table 1); segmented anal-fin rays (modally 17 in A. kenyae 
versus modally 18 in A. mascarenensis and A. straussi; Table 1); 
caudal vertebrae (modally 23 in A. kenyae versus modally 24 in A. 
mascarenensis and A. straussi; Table 3); scales in lateral series (37- 
44, usually 38-41 in A. kenyae versus 4CM-5, usually 41-44 in A. 
mascarenensis and A. straussi; Table 5); anterior lateral-line scales 
(28-35, usually 30-34 in A. kenyae versus 32-39, usually 33-37 in 
A. mascarenensis and A. straussi; Table 6); posterior interorbital 
pores (1-5, usually 2-4 in A. kenyae, 3-4 in A. mascarenensis, and 
1-2 in A. straussi; Table 11); and total parietal pores (usually more 
numerous in A. kenyae and A. mascarenensis than in A. straussi; Fig. 
7). 

The three species are also distinguished on the basis of pre- 
served male coloration. The dorsal and anal fins of A. kenyae and 
A. mascarenensis males are generally pale (mostly red in life), 
with a large dark spot distally on the anterior part of the dorsal fin 
(Figs 4, 9). In contrast, the dorsal and anal fins of A. straussi 
males are generally dusky to black (in life and in preservative), 
with at most an indistinctly darker marking basally on the anterior 
part of the dorsal fin (Fig. 12). Probable intermediate-phase (sex- 
changing) specimens of A. mascarenensis have mostly dusky 
dorsal and anal fins (Fig. 11) and thus might be confused with 
males of A. straussi. However, they bear the characteristic large 
dark spot distally on the anterior part of the dorsal fin. The pelvic 
fins of A. kenyae and A. mascarenensis males are broadly pale on 
the basal part of the fin (though sometimes slightly darker or 
dappled with darker spots on the basalmost portion of the fin) and 
abruptly dark distally, sometimes with a pale distal margin (Figs 
8A-B). In A. straussi males the pelvic fins are mostly dusky, 
although sometimes with a narrow pale basal area, with a pale 
distal margin (Fig. 8C). The bodies of A. kenyae and A. 
mascarenensis males have relatively conspicuous pale spots, 
whereas in A. straussi males pale spots are either absent, or incon- 
spicuous and confined to the anterodorsal part of the body. 

Remarks 

Fourmanoir (1957: 246) recorded an unidentified specimen of 
Anisochromis from Bimbini, Anjouan, Comoros Islands. Based on 
distribution, it would appear that his specimen, which could not be 
located for this study, is referable to A. kenyae. However, there is 



reason to question the generic identity of the specimen. Although 
Fourmanoir gave a dorsal-fin ray count of 1,25, which is character- 
istic of A. kenyae - as perhaps is his count of '35 rangees d'ecailles' 
(= scales in lateral series?) - his counts of anal-fin rays (1,14) and 
lateral-line scales (20) are well outside those known for any species 
of Anisochromis. 

Material Examined 

KENYA: Shimoni, Kisiti Islands, J.L.B. & M.M. Smith, 1 Novem- 
ber 1952, RUSI 854, 14: 17.5-25.0 mm SL (paratypes; 
x-radiographs only), USNM 216415 (out of RUSI 854), 6: 21.7- 

24.5 mm SL (paratypes); Malindi, Sail Rock channel, J.L.B. & 
M.M. Smith, 11 October 1952, RUSI 149, 1: 21.3 mm SL 
(holotype); Shimoni, J.L.B. and M.M. Smith, 27 August 1954, 
RUSI 4905, 3: 22.3-23.9 mm SL. TANZANIA: north-east corner 
of Lathan Island (06°54'05"S 039°55'43'E), rocky shore, Anton 
Bruun Cruise 9, Station HA-6, 20 November 1964, ANSP 134469, 
1: 25.6 mm SL (x-radiograph only). MOZAMBIQUE: Pinda Reef, 
J.L.B. and M.M. Smith, 3 September 1956, RUSI 4906, 7: 21.6- 
25.0 mm SL (1: 23.3 mm SL, subsequently cleared and stained; 2: 
24.0-25.0 mm SL, x-radiographs only). COMOROS ISLANDS: 
Moheli, reef crest off middle of bay at north-east tip of Ouenefou 
Island (12°23'25"S 043°42'330"E), occasional live corals 
(Acropora, Pocillopora and Porites), algae covered rubble, calcar- 
eous rock and occasional sand patches, 0-1 m, R. Winterbottom et 
al., 22 November 1988 (field number RW 88-29), ROM 56502, 9: 
13.8-22.7 mm SL; Anjouan, Point Chongochahari, headland north 
and east of village of M'Jamaoue (12°11'09"S 044°19'03"E), verti- 
cal coral-rock walled gully with sandy floor, small caves and 
crevices, and some live hard corals {Acropora and Pocillopora), 
3-9 m, R. Winterbottom et al., 21 November 1988 (field number 
RW 88-28), ROM 5671 1, 1: 21.6 mm SL; Mayotte, north coast of 
Isle Malandzamiayatsini near eastern tip (12°40'19"S 
044°03'27"E), reef top, profuse soft and hard corals {Acropora, 
Pocillopora, Porites and lettuce coral), with some calcareous rock 
and sand gullies, 0-4 m, W. Holleman et al., 16 November 1988 
(field number RW 88-17), ROM 56501, 2: 22.1-23.0 mm SL. 
MADAGASCAR: Nosy Be, Andilana Beach, 100 m west of hotel, 
50 m offshore (14°43'S 050°57'E), around coral bommies on sand 
flat with turtle grass, 0.3-2.5 m, J. Paxton, B. Collette, D. Cohen, 
E. Anderson, J. Nielsen and K. Sulak, 9 November 1988, AMS 
1.28113-064, 2: 13.9-22.1 mm SL. 

Anisochromis mascarenensis sp.nov. 

Mascarene Annie 

Figs 1,6, 7, 8B, 9-11; Tables 1-11 

Anisochromis kenyae [non Smith, 1954].- Fricke, 1999: 214 
[Reunion]. 

Holotype, SMNS 23037, 23.9 mm SL, Reunion, west coast, Les 
Filaos, L'Hermitage-les-Bains, 11 km south-west of St-Paul, 
21°06'16"S 055°12'38"E, lagoon reef with live corals, 0-0.5 m (low 
tide), R. Fricke, 19 December 1998. 

Paratypes, BMNH 2001 .3.8.2, 23.3 mm SL (subsequently cleared 
and stained), collected with holotype; BPBM 16277, 1: 13.3 mmSL, 
Reunion, Cap Houssaye, sand and coral knoll, 1 2-1 8 m, J.E. Randall, 
27 October 1973; MNHN 2001-494, 24.1 mm SL, collected with 
holotype; SMNS 20933, 2: 19.7-25.5 mm SL, Reunion, west coast, 
Les Filaos, L'Hermitage-les-Bains, 11 km south-west of St-Paul, 
21°06'16"S 055°12'38"E, lagoon reef with dead and live corals, 0- 
0.5 m (at extremely low tide), R.Fricke, 18 December 1998; SMNS 
21025, 4: 19.7-25.2 mm SL, collected with holotype; USNM 364534, 

19.6 mm SL, Mauritius, north coast, Grande Gaube, lagoon reef 






A.C. GILL AND R. FRICKE 




Fig. 6 Distributional records for Anisochromis kenyae (•), A. mascarenensis (■) and A. straussi ( A ). 



with dead and live corals, 3-4 m, P.C. Heemstra, A.C. Gill, M. 
Smale, W. Holleman, P. Clark and B. Galil, 16 May 1995 (field 
number PCH 95-M28). 

Diagnosis 

The following characters distinguish A. mascarenensis from conge- 
ners: dorsal-fin rays 1,25-26, usually 1,26; anal-fin rays 1,17-18, 
usually 1,18; caudal vertebrae 23-24, usually 24; scales in lateral 
series 40-45, usually 42-44; posterior interorbital pores 34; total 
parietal pores 23-44, usually more than 30; dorsal fin of male 
specimens pale in preservative, with conspicuous, large dark spot 
distally on anterior part of fin. 

DESCRIPTION (Based on 1 1 specimens, 13.3-25.5 mm SL) 
Dorsal-fin rays 1,25-26 (1,26), all segmented rays branched; anal-fin 
rays 1,17-18 (1.18), all segmented rays branched; pectoral-fin rays 
13-14 (13/14), upper 1 and lower 0-1 (0/1) rays simple; pelvic-fin 
rays 1,4. medial ray small, inconspicuous and unbranched, all other 
segmented rays branched; upper procurrent caudal-fin rays 5; lower 
procurrent caudal-fin rays 4-5 (4); principal caudal-fin rays 8 + 8, 
upper 0-1 (0) and lower 0-1 (0) unbranched; total caudal-fin rays 
25-26 (25); scales in lateral series 40^5 (43/43); anterior lateral- 



line scales 32-39 (35/35); anterior lateral line terminating beneath 
segmented dorsal-fin ray 19-23 (21/21); predorsal scales 5-6 (5); 
scales in transverse series 10-12 + 1 + 2-3 = 13-16 (11 + 1+2/11 
+ 1 +2); circumpeduncular scales 12; gill rakers 1-2 + 2, sometimes 
with 1-2 tiny rudiments (inconspicuous ossifications bearing a few 
or no teeth) above and below rakers (gill raker count not determined 
in holotype); pseudobranch filaments 6-7 (not determined in 
holotype). 

Cephalic laterosensory pores (all bilaterally paired, unless 
otherwise stated; Fig. 1): nasal pores 1-3 (3/3); anterior interorbital 
pores 2-3 (2/2); posterior interorbital pores (unpaired) ?>-4 (3); 
supraotic pores 2-6 (5/6); suborbital pores 8-13 (10/10); posterior 
otic pores 4-10 (6/5); preopercular pores 12-18 (13/15); dentary 
pores 3-4 (4/4); intertemporal pores 1-2 (1/1); anterior temporal 
pores 1-2 (1/1); posttemporal pores 1-2 (1/1); total parietal pores 
23^4 (35). 

As percentage of standard length (based on nine specimens, 19.6— 
25.5 mm SL): body depth at dorsal-fin origin 18.7-20.8 (19.2); 
greatest body depth 18.7-20.8 (19.2); head width 11.5-13.3(11.7); 
head length 25.2-27.4 (25.9); snout length 4.5-5.6 (5.0); orbit 
diameter 6.7-8.7 (7.1); interorbital width 3.3-4.1 (3.3); upper jaw 



REVISION OF ANISOCHROMINAE 
50 n 



<u 40 
o 



| 3 ° 

Q. 

•s 20 



10- 



10 



15 



20 
mm SL 



25 



30 



Fig. 7. Total counts of parietal pores plotted against SL for specimens of 
Anisochromis kenyae {9), A. mascarenensis (A) and A. straussi (O). 



199 





length 8.6-9.8 (9.6); caudal peduncle depth 12.0-13.7 (13.0); cau- 
dal peduncle length 8.7-10.5 (10.5); predorsal length 28.9-31.1 
(28.9); preanal length 49.6-53.9 (50.2); prepelvic length 21.2-25.5 
(22.6); first segmented dorsal-fin ray length 7.8-10.0 (damaged in 
holotype); third-from-last segmented dorsal-fin ray length 11.6- 
14.5 (13.8); dorsal-fin base length 62.2-66.4 (63.6); first segmented 
anal-fin ray length 7.6-9.8 (8.4); third-from-last segmented anal-fin 
ray length 12.0-14.3 (13.8); anal-fin base length 36.9-41.2 (38.9); 
caudal-fin length 18.3-20.9 (20.1); pectoral-fin length 18.7-21.1 
(20.1); pelvic-fin length 14.3-17.2 (17.2). 

Lower lip complete with deep symphyseal notch; fin spines weak 
and flexible; anterior dorsal-fin pterygiophore formula S/S/(S V ) + 2/ 
1 + 1 (S/S/S v + 2/1 + 1); 21-23 (23) consecutive dorsal-fin 
pterygiophores inserting in 1 : 1 relationship directly behind neural 
spine 4; anterior anal-fin pterygiophore formula 2/1 + 1 or 2 + 1/1 (2/ 

1 + 1); 13-15 (13) consecutive anal-fin pterygiophores inserting in 
1:1 relationship directly behind haemal spine 2; fourth segmented 
pelvic-fin ray longest; caudal fin rounded; ctenoid scales beginning 
at 3-7 (4/3) transverse scale rows behind branchial opening; dorsal 
and anal fins without distinct scale sheaths, though often with 
intermittent scales overlapping fin bases; intermittent series of 
centrally pitted scales originating on midside above anterior part of 
anal fin, extending posteriorly along caudal peduncle to middle part 
of caudal-fin base; additional 1-3 centrally pitted scales present 
above and below pitted scale(s) on middle part of caudal-fin base; 
cheeks, operculum and upper part of head without scales; predorsal 
scales extending anteriorly to point ranging from about 2/3 distance 
from dorsal origin to parietal commissure, to just short of parietal 
commissure (Fig. 1); vertebrae 10 + 23-24 (10 + 24); epurals 2; 
epineurals present on vertebrae 1 through 21-24 (23); pleural ribs 
present on vertebrae 3 through 10, the ultimate rib small to moder- 
ately developed. 

Upper jaw with 3 or 4 (at symphysis) to 1 or 2 (on sides of jaw) 
irregular rows of small conical teeth, those of outer row much larger; 
lower jaw with 2 or 3 (at symphysis) to 1 (on sides of jaw) rows of 
small conical teeth, those of outer row much larger; vomer with 1 or 

2 rows of small conical teeth arranged in chevron; palatines eden- 
tate; tongue edentate and moderately pointed. 

Live coloration 

Males not recorded in detail, but noted to be very similar to that of 
A. kenyae. Females not recorded in detail, but noted to be very 
similar to that of A. kenyae. Probable intermediate phase 
specimens(based on photograph and field notes taken on paratype 
from Mauritius, USNM 364534, 19.6 mm SL, when freshly dead, 
and on field notes taken on paratype from Reunion, SMNS 20933, 




Fig. 8. Ventral view of right pelvic fin of A) Anisochromis kenyae, ROM 
56501, 22.1 mm SL, male; B)A. mascarenensis, SMNS 21025, 25.2 
mm SL. male paratype; C) A. straussi, USNM 216463, 25.2 mm SL, 
male paratype. Scale bars = 0.5 mm. 

25.3 mm SL, when freshly dead): head reddish brown to bright red, 
with black-edged pale pink to white stripe extending from 
posterodorsal rim of orbit, above upper part of preopercle, to upper 
edge of operculum; prominent white streak at posteroventral corner 
of operculum, edged narrowly with black, with prominent black 
spot on mid-anterior edge; large white spot on middle of operculum, 
with scattered smaller white spots on upper edge of preopercle and 
on dorsoposterior part of head; white spots narrowly edged with 
black; two small, white spots on posteroventral rim of orbit, at about 
3 and 5 o'clock positions; narrow dark grey to black bar extending 
from ventral part of orbital rim to posterior edge of maxilla; iris pale 
pink, with radiating dark brown bars; reddish brown coloration 
extending on to upper part of body immediately beneath lateral line, 
grading to dark bluish grey or black elsewhere on body; lateral-line 
scales pale pink; dorsal and anal fins dusky red basally, greyish 
hyaline distally, with white-edged black spot at base of each fin ray, 
anterior part of dorsal fin with large black spot, edged ventrally with 
orange; caudal fin dark grey basally, reddish grey on remainder of 
fin; fleshy pectoral-fin base and base of fin dark grey to black, with 
scattered white spots; remainder of pectoral fin hyaline; pelvic fin 
pale pinkish grey basally, grey to dark grey on remainder of fin, with 
pale grey to hyaline distal margin. 

Preserved coloration 

Males (Fig. 8B, 9) head pale brown, with pale stripe edged with dark 
grey-brown extending from posterodorsal rim of orbit, above upper 
part of preopercle, to upper edge of operculum; narrow brown to 
dark grey-brown bar extending from ventral part of orbital rim to 
posterior edge of maxilla; nape pale brown with dark grey-brown 
bar across parietal commissure; body dark grey-brown to black, 
with scattered indistinct pale brown spots; dorsal fin with large dark 



200 



A.C. GILL AND R. FRICKE 





Fig. 9. Anisochromis mascarenensis, SMNS 23037. 23.9 mm SL, male. 
holotype, Reunion. (Photograph by P. Hurst) 




Fig. 10. Anisochromis mascarenensis, SMNS 21025, 24.2 mm SL. female, 
paratype. Reunion. (Photograph by P. Hurst) 

grey to black spot, extending from first segmented ray to about 
fourth or fifth segmented ray; anal fin and remainder of dorsal fin 
pale brown to white; basal one-quarter of dorsal and anal fins dark 
greyish brown, with small white spot at base of each ray; distal one- 
quarter of dorsal and anal fins greyish hyaline to grey; caudal fin 
dark grey brown to black basally, remainder of fin greyish hyaline; 
pectoral fin dark greyish brown to black with irregular pale brown 
spots basally, remainder of fin greyish hyaline, with fin rays dark 
grey; pelvic fin with broad pale brown band near base of fin, edged 
basally with narrow slightly darker band or dappled spots and 
distally with dark grey (which is darkest immediately adjacent to 
pale band), distal edge of fin pale grey-brown. Females (Fig. 10): 
head brown dorsally, paler ventrally, with two or three dark brown 
bars on nape; large dark brown to dark grey-brown spot on subopercle, 
bordered irregularly with pale brown, sometimes with additional, 
smaller pale-edged dark brown spot on lower part of subopercle; 
brown-edged diffuse pale brown stripe extending from posterodorsal 
rim of orbit, above upper part of preopercle, to upper edge of 
operculum; cheek and operculum sometimes with irregular brown 
vermiform markings and spots; narrow dark brown to grey-brown 
bar extending from ventral part of orbital rim to posterior edge of 
maxilla: body pale brown; upper part of body with brown to grey- 
brown bars, becoming less distinct ventrally; broad pale yellow to 
pale brown area sometimes present on side of body; upper and lower 
edges of caudal peduncle sometimes with small, punctate black 
spots; dorsal and anal fins dusky brown to brownish hyaline, often 
with irregular pale brown small spots: barring on body sometimes 
extending slightly on to fin bases; caudal fin brown to grey brown, 
remainder of fin dusky brown to brownish hyaline; fleshy pectoral- 
fin base brown, with scattered pale spots; pectoral fin brownish 
hyaline to hyaline: pelvic fin dusky brown, pale brown distally. 
Probable intermediate phase specimens (Fig. 1 1 ): pattern generally 
similar to live coloration, reddish brown and red areas on head, body 
and fins become pale brown; white spots and markings on head, 
body and fins less distinct, becoming pale grey to pale yellowish 



Fig. 11. Anisochromis mascarenensis, SMNS 20933. 25.5 mm SL, 

sexually transforming individual(?), paratype. Reunion. (Photograph by 
P. Hurst) 

brown; dark spots and markings on head, body and fins become dark 
brown to dark grey-brown; indistinct dark spot or ocellated dark spot 
(as in females) variably present on subopercle; body either more-or- 
less uniformly dark grey-brown (except for pale spots) or dark 
grey-brown with narrow pale brown bands; pelvic fins mostly dusky 
brown to dark grey-brown, with broad pale band near fin base and 
pale brown to hyaline distal margin. 

Habitat and Distribution 

Anisochromis mascarenensis is known only from Reunion and 
Mauritius, Mascarene Islands (Fig. 6). It has been collected from 
lagoon reefs with live and dead corals in 0-18 m. Most specimens 
collected by the second author at Reunion emerged from the base of 
live, branched Acropora. 

Comparisons with other species 
See under A. kenyae. 

Remarks 

Anisochromis mascarenensis is apparently not common in Mauri- 
tius. The first author and associates made 13 rotenone stations in 
apparently appropriate habitat (around shallow lagoonal reefs) in 
Mauritius, but collected only a single specimen. 

Etymology 

The specific epithet alludes to the distribution of the species. 

Material Examined 

See above under type material. 

Anisochromis straussi Springer, Smith & Fraser, 1977 

Saint Brandon's Annie 

Figs 6, 7, 8C, 12-13; Tables 1-11 

Anisochromis straussi Springer, Smith & Fraser, 1977: 2, figs la, lb 
and 2 [type locality: 2 miles east of Raphael Island, Saint Brandon's 
Shoals; holotype USNM21642].-Godkin& Winterbottom, 1985: 
634, fig. ID [osteological and myological comparison]- Mooi, 
1990: 457, tables 1,3, fig. 2e [egg surface morphology]. 

Diagnosis 

The following characters distinguish A. straussi from congeners: 
dorsal-fin rays 1,25-27, usually 1,26; anal-fin rays 1,17-19, usually 
1,18; caudal vertebrae 23-25, usually 24; scales in lateral series 41- 
45, usually 41-44; anterior lateral-line scales 32-39, usually 33-37; 
posterior interorbital pores 1-2; total parietal pores 14-30; and 
dorsal fin of male specimens generally dusky in preservative, with- 
out large dark spot on anterior part of fin. 

Description (Based on 82 specimens, 16.1-28.3 mm SL) 
Dorsal-fin rays 1,25-27 (1,26), all segmented rays branched; anal-fin 
rays 1,17-19 (1,18), all segmented rays branched; pectoral-fin rays 



REVISION OF ANISOCHROMINAE 



201 



13-15 (14/14), upper 1-2 (1/1) and lower 0-1 (0/0) rays simple; 
pelvic-fin rays 1,4, medial ray small, inconspicuous and unbranched, 
all other segmented rays branched; upper procurrent caudal-fin rays 
4-6 (5); lower procurrent caudal-fin rays 3-4 (4); principal caudal- 
fin rays 8 + 8, upper 0-1 (0) and lower 0-1 (0) unbranched; total 
caudal-fin rays 23-26 (25); scales in lateral series 41^15 (43/43); 
anterior lateral-line scales 32-39 (32/34); anterior lateral line ter- 
minating beneath segmented dorsal-fin ray 1 8-24 (18/19); predorsal 
scales 4—7 (6); scales in transverse series 1 1-14 + 1 + 2-3 = 14-18 
(12 + 1 + 3/13 + 1+2); circumpeduncular scales 12-14 (12); gill 
rakers 2 + 2-3, sometimes with 1-2 tiny rudiments (inconspicuous 
ossifications bearing a few or no teeth) above and below rakers (gill 
raker count not determined in holotype); pseudobranch filaments 6- 
7 (not determined in holotype). 

Cephalic laterosensory pores (all bilaterally paired, unless 
otherwise stated): nasal pores 2-3 (2/2); anterior interorbital pores 
1-3 (2/2); posterior interorbital pores (unpaired) 1-2 (2); supraotic 
pores 1-3 (3/2); suborbital pores 8-13 (12/11); posterior otic pores 
2-7 (3/2); preopercular pores 9-15 (13/13); dentary pores 3^4 (4/4); 
intertemporal pores 1-2 (1/1); anterior temporal pores 0-1 (1/1); 
posttemporal pores 1; total parietal pores 14-30 (25). 

As percentage of standard length (based on 20 specimens, 16.7- 
27.0 mm SL): body depth at dorsal-fin origin 19.2-21.8 (21.8); 
greatest body depth 20.6-23.8 (23.8); head width 11.5-13.7 (12.1); 
head length 24.6-28.6 (27.2); snout length 4.2-5.7 (5.4); orbit 
diameter 6.5-9.0 (7.1); interorbital width 3.3^4.5 (3.8); upper jaw 
length 9.1-10.5 (10.5); caudal peduncle depth 12.7-15.6 (14.2); 
caudal peduncle length 8.2-10.7 (9.6); predorsal length 27.7-31.7 
(30.5); preanal length 48.9-52.9 (49.8); prepelvic length 22.6-25.7 
(23.4); first segmented dorsal-fin ray length 7.9-10.1 (7.9); third- 
from-last segmented dorsal-fin ray length 1 1 .7-1 4.9 ( 14.2); dorsal-fin 
base length 62.0-66.9 (66.9); first segmented anal-fin ray length 
6.9-8.9 (7.1); third-from-last segmented anal-fin ray length 12.3— 
14.4 (13.8); anal-fin base length 38. 1-41.8 (41 .0); caudal-fin length 
17.8-20.7 (20.5); pectoral-fin length 15.9-20.8 (19.2); pelvic-fin 
length 10.5-17.8(15.1). 

Lower lip complete with deep symphyseal notch; fin spines weak 
and flexible; anterior dorsal-fin pterygiophore formula S/S/(S V ) + 2/1 
+ 1 (S/S/2/1 + 1); 21-24 (23) consecutive dorsal-fin pterygiophores 
inserting in 1 : 1 relationship directly behind neural spine 4; anterior 
anal-fin pterygiophore formula 2/1 + 1,2+ 1/1 or 2+ 1 + 1/1 (2/1 + 1 ); 
12-16 (13) consecutive anal-fin pterygiophores inserting in 1:1 
relationship directly behind haemal spine 2; fourth segmented pelvic- 
fin ray longest; caudal fin rounded; ctenoid scales beginning at 3-14 
(4/3) transverse scale rows behind branchial opening; dorsal and anal 
fins without distinct scale sheaths, though often with intermittent 
scales overlapping fin bases; intermittent series of centrally pitted 
scales originating on midside above anterior part of anal fin, extending 
posteriorly along caudal peduncle to middle part of caudal-fin base; 
additional 1-3 centrally pitted scales present above and below pitted 
scale(s) on middle part of caudal-fin base; cheeks, operculum and 
upper part of head without scales ; predorsal scales extending anteriorly 
to point ranging from about 2/3 distance from dorsal origin to parietal 
commissure, to just short of parietal commissure; vertebrae 10-1 1 + 
23-25 = 33-35 ( 10 + 24); epurals 2; epineurals present on vertebrae 1 
through 20-27 (22) ; pleural ribs present on vertebrae 3 through 1 0, the 
ultimate rib very small to moderately developed. 

Upper jaw with 3 or 4 (at symphysis) to 1 or 2 (on sides of jaw) 
irregular rows of small conical teeth, those of outer row much larger; 
lower jaw with 2 or 3 (at symphysis) to 1 (on sides of jaw) rows of 
small conical teeth, those of outer row much larger; vomer with 1 or 
2 rows of small conical teeth arranged in chevron; palatines eden- 
tate; tongue edentate and moderately pointed. 



Live coloration 

Males (based on a photograph of a probable paratype, and on the 
description given by Springer et al., 1977: 4) head bright reddish 
orange, with black-edged white stripe extending from posterodorsal 
rim of orbit, above upper part of preopercle, to upper edge of 
operculum; two white spots on posteroventral rim of orbit, at about 
3 and 5 o'clock positions; narrow dark grey bar extending from 
ventral part of orbital rim to posterior edge of maxilla; iris yellow, 
red centrally, with radiating brown bars; reddish orange coloration 
extending slightly on to anterior part of body, rapidly grading to 
uniform black; dorsal, anal and caudal fins black basally, becoming 
grey to greyish hyaline on distal margin; base of dorsal fin with 
small, intermittent pale grey spots; pectoral fin dark grey to black 
basally, remainder of fin greyish hyaline, with fin rays dark grey; 
pelvic fin black, with distal margin pale grey to hyaline. Females 
(based on photographs of two probable paratypes, and on the 
description given by Springer et al., \911: 4) head olive-brown to 
brown dorsally, becoming pale green to lime green ventrally, with 
two pale olive bars on nape; large dark grey to black spot on 
subopercle, bordered irregularly with white; black-edged white 
stripe extending from posterodorsal rim of orbit, above upper part of 
preopercle, to upper edge of operculum; two white spots or clusters 
of white spots on posteroventral rim of orbit, at about 3 and 5 o'clock 
positions; narrow dark brown to grey bar extending from ventral part 
of orbital rim to posterior edge of maxilla; iris red, with radiating 
brown bars; body generally orange-brown, becoming olive to dusky 
green posteriorly; dorsal part of body with short, indistinct olive- 
brown bars, alternating with pale olive to orange-brown bars; pale 
olive to orange-brown bars sometimes extending on to lower part of 
body, becoming pale pink ventrally; broad orange-yellow area often 
present on side of body; dorsal and anal fins dusky green to dusky 
orange; barring on upper part of body extending slightly on to 
dorsal-fin base; dark bars on posterior part of dorsal-fin base some- 
times bearing dark grey to black punctate spots; distal margins of 
dorsal and anal fins abruptly pale grey to hyaline; caudal fin olive to 
lime green basally, remainder of fin dusky olive to dusky orange; 
fleshy pectoral-fin base dusky orange to dusky olive, with scattered 
small white spots; pectoral fin lime green basally, becoming green- 
ish to orangish hyaline distally; pelvic fin olive to grey basally, 
remainder of fin dark olive to dark grey, with pale grey to hyaline 
distal margin. 

Preserved coloration 

Males (Fig. 8C, 12): pattern generally similar to live coloration, 
head and anterior part of body becoming pale brown, paler ventrally; 
pale markings on head obsolete; dark spots and stripes on head 
remain, though sometimes faint, becoming brown to dark grey- 
brown; body behind pectoral-fin base dark grey-brown; several 
indistinct pale brown spots sometimes present on anterodorsal part 




Fig. 12. Anisochromis straussi, USNM 216463, 24.7 mm SL, male, 
paratype, Saint Brandon's Shoals. (Photograph by P. Hurst) 



202 



A.C. GILL AND R. FRICKE 




Fig. 13. Anisochromis straussi, USNM 216463. 22.8 mm SL. female, 
paiatype, Saint Brandon's Shoals. (Photograph by R Hurst) 

of body: coloration of fins similar to when live; pale brown spots 
often present at base of each dorsal-fin ray, several pale brown spots 
sometimes present basally on anterior part of dorsal fin. Females 
(Fig. 13): pattern generally similar to live coloration, head and body 
becoming pale brown; pale markings on head and body indistinct or 
absent; dark spots and stripes on head mostly remain, becoming dark 
grey-brown to dark brown (ocellated spots sometimes absent, though 
possibly these represent intermediately coloured, sex-transforming 
specimens), sometimes with additional irregular brown spots and 
markings on cheek and operculum; dark barring on head and body 
variably remains, becoming brown to pale brown; dark punctate 
spots on caudal peduncle remain, becoming dark brown to dark 
grey-brown, sometimes extending anteriorly to near middle of 
dorsal- and anal-fin bases, ocassionally extending on to sides of 
body; dorsal and anal fins dusky brown to brownish hyaline, often 
with irregular pale brown small spots; barring on body sometimes 
extending slightly on to fin bases; caudal fin pale brown, remainder 
of fin pale brown to hyaline; pectoral-fin base pale brown, remain- 
der of fin pale brown to hyaline; pelvic fin dusky brown to brownish 
hyaline, paler on base and distal margin. 

Habitat and Distribution 

Anisochromis straussi is known only from Saint Brandon's Shoals 
(Fig. 6). According to Springer et al. (1977: 6), it was collected in 
0.25-1 1 m from rocky reefs that included dead and live coral, 
proximate to areas exposed at low tide. They further noted that 
'specimens of A. straussi were lying on the bottom adjacent to 
isolated, small (perhaps less than 0.5 meter in diameter), live coral 
heads with surfaces composed of tiny finger-like projections. Our 
presumption is that the Anisochromis were living in the corals.' 

Comparisons with other species 
See under A. kenyae. 

Remarks 

Springer et al. ( 1977) gave a standard length of 25.5 mm for the 

holotype of A. straussi, whereas we measured it as only 23.9 mm. 

Material Examined 

Saint Brandon's Shoals (= Cargados Carajos) Lagoon at Tortue 
Island (16°19'S 059°41'E), 0.15 m (stated depth 0.5 feet), V.G. 
Springer et al., 7 April 1976 (field number VGS 76-11), USNM 
216463, 19: 16.2-27.0 mm SL (paratypes); 2 miles east of Raphael 
Island (16°20'S 059°38.5'E), inside edge of reef flat, 0.15-1.05 m 
(stated depth 0.5-3.5 feet), V.G. Springer et al, 3 April 1976 (field 
number VGS 76-7), USNM 216462, 1: 23.9 mm SL (holotype), 
USNM 215859, 26: 18.7-26.1 mm SL (paratypes; x-radiographs 
only); off northern tip of Saint Brandon's Shoals, ca. 16°25'S 
59°36'E, rocky reef with some live coral and some channels and 
white coarse sand bottom, 6-10.5 m (stated depth 20-35 feet), V.G. 
Springer et al., 6 April 1976 (field number VGS 76-10), BMNH 



1976.8.24.1-10, 10: 16.8-25.4 mm SL (paratypes; 21.5 mm SL 
paratype subsequently cleared and stained); about 100 yards off 
west side of Raphael Island (ca. 16°26'S 059°36'E), coral patch in 
surge channel, 0-7.5 m (stated depth 0-25 feet), V.G. Springer etal, 
2 April 1976 (field number VGS 76-6), USNM 216464, 1: 26.8 mm 
SL (paratype; x-radiograph only); lagoon south of Raphael Island 
(ca. 16°28'S 059°37'E), live and dead coral reef surrounded by fine 
white sand, 0-3.6 m (stated depth 0-12 feet), V.G. Springer et al., 8 
April 1 976 (field number VGS 76- 1 2), USNM 2 1 6466, 2: 26.5-28.3 
mm SL (paratypes: x-radiographs only); along southeast side of 
Grande Passe (ca. 16°28'S 059°40'E), face and channels of reef, 0- 
6 m (stated depth 0-20 feet), V.G. Springer et al., 5 April 1976 (field 
number VGS 76-9), CAS 37640, 14: 16.1-24.9 mm SL (paratypes; 
x-radiographs only); ca. 16°32'S 059°41'E, 0-2 m, V.G. Springer et 
al., 30 March 1976 (field number VGS 76-1), USNM 216465, 3: 
22.6-23.9 mm SL (paratypes; x-radiographs only); ca. 16°43'S 
059°35'E, live coral reef with dead rock and coral, rubble shore, 0- 
1.2 m (stated depth 0-4 feet), V.G. Springer et al, 11 April 1976 
(field number VGS 76-17), AMNH 35892, 6: 22.0-28.0 mm SL 
(paratypes; x-radiographs only). 



KEY TO SPECIES OF ANISOCHROMIS 



la. Dorsal-fin rays 1,25-26, usually 1,25; anal-fin rays 1,17-18, usually 
1,17; caudal vertebrae 22-24, usually 23; scales in lateral series 31-44, 
usually 38 — 4 1 ; anterior lateral-line scales 28-35, usually 30-34 (east 
Africa, Comoros Islands and Madagascar) kenyae Smith 

lb Dorsal-fin rays 1,25-27, usually 1,26; anal-fin rays 1,17-19, usually 
1,18: caudal vertebrae 23-25, usually 24; scales in lateral series 40-45, 
usually 41^44-; anterior lateral-line scales 32-39. usually 33-37 2 

2a Dorsal fin of male specimens pale in preservative, with conspicuous, 
large dark spot distally on anterior part of fin; pelvic fins of preserved 
males broadly pale on basal part of fin (sometimes slightly darker or 
dappled with darker spots on basalmost portion of fin) and abruptly dark 
distally, sometimes with pale distal margin: posterior interorbital pores 
3^1; total parietal pores 23—44, usually more than 30 (Reunion and 
Mauritius) mascarenensis sp.nov. 

2b Dorsal fin of male specimens generally dusky in preservative, without 
large dark spot on anterior part of fin; pelvic fins of males mostly dusky 
in preservative, although sometimes with narrow pale basal area, with 
pale distal margin; posterior interorbital pores 1-2; total parietal pores 
14—30 (Saint Brandon's Shoals) straussi Springer, Smith & Fraser 



ACKNOWLEDGEMENTS. We thank the following variously for assistance 
with radiography or photography or for the loan of specimens, radiographs or 
photographs: A. Bentley, O.A. Crimmen, S. Davidson, J. P. Garcia, P.C. 
Heemstra. A.-M. Hine, P. Hurst, S.L. Jewett. M. McGrouther, L. Palmer, J.E. 
Randall. S.J. Raredon, S.E. Reader. V.G. Springer. A. Suzumoto, T. Trnski 
and R. Winterbottom. The first author thanks the other members of the 1995 
Mauritius expedition for assistance in the field: P. Clark. B. Galil, P.C. 
Heemstra, W. Holleman, M.J. Smale and D.G. Smith. The success of the 
expedition also owes much to the kind assistance of D. Pelicier and of 
Mauritian Fisheries officials, particularly staff of the Albion Fisheries Research 
Centre. E. de Chavanes (Directeur, Directoire Regionale et Departementale 
des Affaires Maritimes, Saint-Denis, Reunion) granted a fish collecting and 
export permit to the second author. The second author is grateful to the 
German Research Council (DFG) for financial support of research trips to the 
Mascarenes in 1995 and 1998/1999. We thank V.G. Springer for helpful 
discussions and access to materials, and R.D. Mooi and R. Winterbottom for 
critically reviewing the manuscript and providing helpful suggestions. 



REVISION OF ANISOCHROMINAE 



203 



REFERENCES 



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(Pisces, Perciformes) of the eastern Pacific: kinds, distributions, and early life 
histories and observations of five of these from the northwest Atlantic. Bulletin of 
Marine Science 26(3): 285^02. 

Bailey, R.M., Robins, C.R., & Greenwood, P.H. 1980. Chromis Cuvier in Desmarest, 
1814 (Osteichthyes, Perciformes, Pomacentridae): proposal to place on official list 
of generic names in zoology, and that the generic names ending in -chromis be ruled 
to be masculine. Z.N.(S.) 2329. Bulletin of Zoological Nomenclature 37(4): 247- 
255. 

Fourmanoir, P. 1957. Poissons teleosteens des eaux Malgaches du Canal de Mozam- 
bique. Memoires de Vlnstitut Scientifique de Madagascar, ser. F, 1: 1-316, pis 1—17. 

Fricke, R. 1999. Fishes of the Mascarene Islands (Reunion, Mauritius, Rodriguez). An 
annotated checklist, with descriptions of new species. Theses Zoologicae 31: 1-759. 

Fujita, K. 1989. Nomenclature of cartilaginous elements of the caudal skeleton in 
teleostean fishes. Japanese Journal of Ichthyology 36(1): 22-29. 

Gill, A.C. 1998. Homology of the anterior vertebrae, ribs, and dorsal fin pterygiophores 
and rays in congrogadine fishes (Perciformes: Pseudochromidae). Copeia 1998(4): 
1041-1045. 

In press. Revision of the Indo-Pacific fish subfamily Pseudochrominae 

(Perciformes: Pseudochromidae). Ichthyological Monographs of the J.L.B. Smith 
Institute of Ichthyology 3. 

& Edwards, A.J. 1999. Monophyly. interrelationships and description of three 

new genera in the dotty back fish subfamily Pseudoplesiopinae (Teleostei: Perciformes: 
Pseudochromidae). Records of the Australian Museum 51(2): 141-160. 

& . In press. Revision of the Indian Ocean dottyback fish genera Chlidichthys 

and Pectinochromis (Perciformes: Pseudochromidae: Pseudoplesiopinae). Ichthyo- 
logical Bulletin of the J.L.B. Smith Institute of Ichthyology 

, Mooi, R.D. & I Interims. J.B. 2000. Description of a new subgenus and species 

of the fish genus Congrogadus Gunther from Western Australia (Perciformes: 
Pseudochromidae). Records of the Western Australian Museum 20(1): 69-79. 

Godkin, C.M., & Winter-bottom, R. 1985. Phylogeny of the family Congrogadidae 
(Pisces; Perciformes) and its placement as a subfamily of the Pseudochromidae. 
Bulletin of Marine Science 36(3): 633-671. 



Greenwood, P.H. 1995. Preliminary studies on a mandibulohyoid 'ligament' and other 

intrabuccal connective tissue linkages in cirrhitid, latrid and cheilodactylid fishes 

(Perciformes: Cirrhitoidei). Bulletin of the Natural History Museum, London, Zool- 
ogy Series 61(2): 91-101. 
International Commission on Zoological Nomenclature. 1986. Opinion 1417. 

Chromis Cuvier in Desmarest, 1814 (Osteichthyes, Perciformes): gender confirmed 

as feminine. Bulletin of Zoological Nomenclature 43(3): 267-268. 
Johnson, G.D., & Patterson, C. 1993. Percomorph phylogeny: a survey of 

acanthomorphs and a new proposal. Bulletin of Marine Science 52(1): 554-626. 
Leviton, A.E., Gibbs, R.H., Jr, Heal, E., & Dawson, C.E. 1985. Standards in 

herpetology and ichthyology. Part 1. Standard symbolic codes for institutional 

resource collections in herpetology and ichthyology. Copeia 1985(3): 802-832. 
McAllister, D.E. 1968. Evolution of branchiostegals and classification of teleostome 

fishes. National Museum of Canada Bulletin, no. 221: 1-239. 
Mooi, R.D. 1990. Egg surface morphology of pseudochromoids (Perciformes: 

Percoidei), with comments on its phylogenetic implications. Copeia 1990(2): 455- 

475. 
Smith, J.L.B. 1954. The Anisochromidae: a new family of fishes from east Africa. 

Annals and Magazine of Natural History, ser. 12, 7: 298-302, pi. 6. 
Smith, M.M. 1977. A note on Anisochromis kenyae. Ichthyological Bulletin of the 

J.L.B. Smith Institute of Ichthyology, no. 35: 22-23, pi. 4C-B. 
1986. Family no. 169: Pseudochromidae. pp. 539-541. In: Smith, M.M. & 

Heemstra, PC. (eds) Smiths' Sea Fishes. Macmillan South Africa, Johannesburg. 
Springer, V.G., Smith, C.L., & Fraser, T.H. 1 977. Anisochromis straussi, new species 

of protogynous hermaphroditic fish, and synonymy of Anisochromidae. 

Pseudoplesiopidae. and Pseudochromidae. Smithsonian Contributions to Zoology. 

no. 252: 1-15. 
Taylor, W.R. & Van Dyke, G.C. 1985. Revised procedures for staining and clearing 

small fishes and other vertebrates for bone and cartilage study. Cybium 9(2): 107-1 19. 
Wheeler, A. 1985. The World Encyclopedia of Fishes. Macdonald & Co, London. 
Winterbottom, R. 1986. Revision and vicariance biogeography of the subfamily 

Congrogadinae (Pisces: Perciformes: Pseudochromidae). Indo-Pacific Fishes, no. 9: 

1-34, pi. 1. [Dated 1985, but actually published 24 February 1986] 
1996. A new species of the congrogadin genus Rusichthys from southern Oman 

(Perciformes: Pseudochromidae). with notes on its osteology. Canadian Journal of 

Zoology 74(3): 581-584. 



Table 1 Frequency distributions of dorsal-, anal- and pectoral-fin-ray counts for Anisochromis species. Bilateral counts of pectoral-fin rays are included. 



25 



Segmented D rays 
26 27 x 



SD 



17 



Segmented A rays 
18 19 x 



SD 



13 



Pectoral rays 
14 15 x 



SD 



A. kenyae 

Kenya 

Tanzania 

Mozambique 

Comoros 

Madagascar 

Total 
A. mascarenensis 

Reunion 

Mauritius 

Total 
A. straussi 



22 


2 


7 


_ 


9 


3 


39 


6 


1 


9 


1 


10 


10 


69 



25. 



0.3 



23 



25.0 


0.0 


7 


- 


25.3 


0.5 


10 


2 


25.1 


0.3 


42 


4 


25.9 


0.3 


2 


8 






- 


1 


25.9 


0.3 


2 


9 


25.9 


0.4 


5 


74 



17.0 


0.2 


1 


15 
Mo data taken 


4 


14.2 


0.5 


17.0 


0.0 


1 


9 


- 


13.9 


0.3 


17.2 


0.4 


4 


20 


- 


13.8 


0.4 


17.5 


0.7 


- 


4 


- 


14.0 


0.0 


17.1 


0.3 


6 


48 


4 


14.0 


0.4 


17.8 


0.4 


7 


13 


_ 


13.7 


0.5 






2 


- 


- 


13.0 


0.0 


17.8 


0.4 


9 


13 


- 


13.6 


0.5 


18.0 


0.3 


8 


51 


1 


13.9 


0.4 



204 



A.C. GILL AND R. FRICKE 



Table 2 Frequency distributions of caudal-fin-ray counts for Anisochromis species. 



Upper 
6 



Procurrent caudal-fin rays 
SD 3 



Lower 

5 



SD 



A. kenyae 
Kenya 

Tanzania 

Mozambique 

C omoros 

Madagascar 

Total 
A. mascarenensis 

Reunion 

Mauritius 

Total 
A. straussi 



A. kenyae 

Kenya 

Tanzania 

Mozambique 

Comoros 

Madagascar 

Total 
A. mascarenensis 

Reunion 

Mauritius 

Total 
A. straussi 



3 

23 



20 

1 
5 

10 
2 

38 

10 

1 

11 

74 



24 

1 



25 

15 
1 

4 
8 
2 

30 



1 

9 
72 



5.0 

5.2 
5.2 
5.0 
5.1 

5.0 

5.0 
5.0 

26 

6 

2 
2 

10 

2 

2 
? 



0.4 

0.4 
0.4 
0.0 
0.4 

0.0 

0.0 
0.3 



1 



Total caudal-fin rays 



27 

1 

2 
3 



25.3 

25.3 
25.5 
25.0 
25.3 

25.2 

25.2 
25.0 



17 
1 

6 
8 
2 

34 



1 
9 

78 



SD 

0.6 

0.5 
0.8 
0.0 
0.6 

0.4 

0.4 
0.3 



1 
4 

11 

2 

2 



4.3 

4.1 
4.3 
4.0 
4.2 

4.2 

4.2 
4.0 



0.4 

0.4 
0.5 
0.0 
0.4 

0.4 

0.4 
0.1 



Table 3 Frequency distributions of vertebral counts for Anisochromis species. 



10 



Precaudal 



11 



SD 



22 



23 



Caudal 

24 25 



SD 



. kenyae 

Kenya 

Tanzania 

Mozambique 

Comoros 

Madagascar 

Total 
. mascarenensis 

Reunion 

Mauritius 

Total 
. straussi 



A. kenyae 

Kenya 

Tanzania 

Mozambique 

Comoros 

Madagascar 

Total 
A. mascarenensis 

Reunion 

Mauritius 

Total 
A. straussi 



24 

1 
7 

12 
2 

46 

10 

1 

11 

79 

32 

2 

1 
1 

4 



33 

21 
1 
6 

II 
I 

40 

2 

2 
10 



10.0 



10.0 

10.0 
10.0 

34 



8 
1 

9 
70 



0.0 



10.0 


0.0 


10.0 


0.0 


10.0 


0.0 


10.0 


0.0 



0.0 

0.0 
0.2 



Total 



35 



33.0 



33.8 

33.8 
33.9 



21 


1 


1 


- 


6 

11 


- 


1 
40 


1 
2 


2 


8 


- 


1 


2 


9 


12 


68 



SD 



0.4 



32.9 


0.4 


32.9 


0.3 


33.5 


0.7 


33.0 


0.4 



0.4 

0.4 
0.4 



23.0 



23.8 

23.8 
23.9 



0.4 



22.9 


0.4 


22.9 


0.3 


23.5 


0.7 


23.0 


0.4 



0.4 

0.4 
0.4 



REVISION OF ANISOCHROMINAE 



205 



Table 4 Frequency distributions of numbers of consecutive dorsal-fin pterygiophores inserting in 1:1 relationship with interneural spaces directly behind 
neural spine 4, and anal-fin pterygiophores inserting in 1 : 1 relationship with interhaemal spaces directly behind haemal spine 2 for species of 
Anisochromis. 









1:1 D 


pterygiopr 


lores 










1:1 A 


pterygioph 


tores 








20 


21 


22 


23 


24 


X 


SD 


12 


13 


14 


75 


16 


X 


SD 


A. kenyae 






























Kenya 


1 


1 


21 


1 


- 


21.9 


0.5 


1 


- 


22 


1 


- 


14.0 


0.5 


Tanzania 


- 


- 


1 


- 


- 






- 


- 


1 


- 


- 






Mozambique 


- 


- 


7 


- 


- 


22.0 


0.0 


1 


- 


6 


- 


- 


13.7 


0.8 


Comoros 


1 


2 


7 


- 


- 


21.6 


0.7 


3 


2 


4 


- 


- 


13.1 


0.9 


Madagascar 


- 


- 


1 


- 


- 






- 


- 


1 


1 


- 


14.5 


0.7 


Total 


2 


3 


37 


1 


- 


21.9 


0.5 


5 


2 


34 


2 


- 


13.8 


0.7 


A. mascarenensis 






























Reunion 


- 


2 


2 


5 


- 


22.3 


0.9 


- 


3 


2 


4 


- 


14.1 


0.9 


Mauritius 


- 


- 


1 


- 


- 






- 


- 


- 


1 


- 






Total 


- 


2 


3 


5 


- 


22.3 


0.8 


- 


3 


2 


5 


- 


14.2 


0.9 


A. straussi 


- 


14 


8 


57 


2 


22.6 


0.8 


1 


23 


6 


48 


1 


14.3 


1.0 



Table 5 Frequency distributions of counts of scales in lateral series for Anisochromis species. Bilateral counts are included. 





37 


38 


39 


40 


41 


42 


43 


44 


45 


X 


SD 


A. kenyae 
























Kenya 


- 


2 


5 


5 


4 


2 


1 


1 


- 


40.3 


1.6 


Mozambique 


- 


- 


3 


3 


3 


1 


- 


- 


- 


40.2 


1.0 


Comoros 


1 


4 


7 


8 


4 


- 


- 


- 


- 


39.4 


1.1 


Madagascar 


- 


- 


2 


1 


1 


- 


- 


- 


- 


39.8 


1.0 


Total 


1 


6 


17 


17 


12 


3 


1 


1 


- 


39.9 


1.3 


A. mascarenensis 
























Reunion 


- 


- 


- 


1 


- 


5 


4 


5 


2 


43.1 


1.3 


Mauritius 


- 


- 


- 


- 


- 


1 


- 


1 


- 


43.0 


1.4 


Total 


- 


- 


- 


1 


- 


6 


4 


6 


2 


43.1 


1.3 


A. straussi 


- 


- 


- 


- 


6 


19 


20 


13 


1 


42.7 


1.0 



Table 6 Frequency distributions of counts of anterior lateral-line scales for Anisochromis species. Bilateral counts are included. 





28 


29 


30 


31 


32 


33 


34 


35 


36 


37 


38 


39 


X 


SD 


A. kenyae 






























Kenya 


1 


1 


3 


4 


1 


- 


3 


1 


- 


- 


- 


- 


31.4 


2.1 


Mozambique 


- 


- 


1 


2 


1 


4 


1 


- 


- 


- 


- 


- 


32.2 


1.3 


Comoros 


3 


2 


4 


5 


4 


3 


- 


- 


- 


- 


- 


- 


30.7 


1.6 


Madagascar 


- 


- 


1 


2 


1 


- 


- 


- 


- 


- 


- 


- 


31.0 


0.8 


Total 


4 


3 


9 


13 


7 


7 


4 


1 


- 


- 


- 


- 


31.2 


1.7 


A. mascarenensis 






























Reunion 


- 


- 


- 


- 


1 


2 


2 


7 


3 


1 


1 


1 


35.2 


1.7 


Mauritius 


- 


- 


- 


- 


- 


- 


- 


1 


1 


- 


- 


- 


35.5 


0.7 


Total 


- 


- 


- 


- 


1 


2 


2 


8 


4 


1 


1 


1 


35.2 


1.6 


A. straussi 


- 


- 


- 


- 


2 


7 


13 


12 


10 


6 


3 


1 


35.0 


1.6 



Table 7 Frequency distributions of anterior lateral-line termination positions (relative to segmented dorsal-fin rays) for Anisochromis species. Bilateral 
counts are included. 



17 



18 



19 



20 



Segmented dorsal-fin ray 
21 22 



23 



24 



SD 



A. kenyae 
















Kenya 


1 


4 


4 


3 


2 


- 


- 


Mozambique 


- 


1 


3 


3 


2 


- 


- 


Comoros 


2 


3 


9 


3 


4 


- 


- 


Madagascar 


- 


- 


- 


2 


- 


- 


- 


Total 


3 


8 


16 


11 


8 


- 


- 


A. mascarenensis 
















Reunion 


- 


- 


1 


5 


6 


4 


2 


Mauritius 


- 


- 


- 


- 


1 


1 


- 


Total 


- 


- 


1 


5 


7 


5 


2 


A. straussi 


- 


1 


3 


9 


19 


12 


9 



19.1 


1.2 


19.7 


1.0 


19.2 


1.2 


20.0 


0.0 


19.3 


1.1 


21.1 


1.1 


21.5 


0.7 


21.1 


1.1 


21.3 


1.3 



206 



A.C. GILL AND R. FRICKE 



Table 8 Frequency distributions of counts of scales in transverse series for Anisochromis species. Bilateral counts are included. 



10 



II 



Scales below lateral line 
12 13 14 



SD 



Scales above lateral line 
3 x 



SD 



A. kenyae 

Ken\a 

Mozambique 
Comoros 
Madagascar 
Total 
.4. mascarenensis 
Reunion 
Mauritius 



7 

4 

5 

2 

18 



7 

6 

13 

1 

27 



6 

1 
11 



Total 


1 


12 


7 


A. straussi 


- 


15 


33 




13 


14 


15 


A. kenyae 








Kenya 

Mozambique 

Comoros 


1 
4 
5 


1 

5 

13 


4 
1 
6 


Madagascar 
Total 


2 
18 


1 
26 


1 

12 


A. mascarenensis 








Reunion 


- 


11 


6 


Mauritius 


1 


1 


- 


Total 


1 


12 


6 


A. straussi 


- 


12 


22 



10.9 


0.9 


19 


1 


2.1 


0.2 


10.6 


0.5 


9 


1 


2.1 


0.3 


11.0 


0.7 


24 


- 


2.0 


0.0 


10.8 


1.0 


4 


- 


2.0 


0.0 


10.9 


0.8 


56 


2 


2.0 


0.2 


11.4 


0.5 


17 


1 


2.1 


0.2 


10.5 


0.7 


2 


- 


2.0 


0.0 


11.3 


0.6 


19 


1 


2.1 


0.2 


11.9 


0.7 


36 


22 


2.4 


0.5 



Total scales in transverse series 
16 17 18 



19 



SD 



14.0 


1.1 


13.7 


0.7 


14.0 


0.7 


13.8 


1.0 


13.9 


0.8 


14.4 


0.6 


13.5 


0.7 


14.4 


0.7 


15.3 


0.9 



Table 9 Frequency distributions of counts of predorsal and circumpeduncular scales for Anisochromis species. 



Predorsal scales 
6 7 



SD 



12 



Circumpeduncular scales 
13 14 x 



SD 



A. kenyae 










Kenya 


- 


1 


5 


3 


Mozambique 


1 


1 


1 


1 


Comoros 


2 


6 


4 


- 


Madagascar 


- 


2 


- 


- 


Total 


3 


10 


10 


4 


A. mascarenensis 










Reunion 


- 


5 


4 


- 


Mauritius 


- 


1 


- 


- 


Total 


- 


6 


4 


- 


A. straussi 


5 


13 


11 


1 



6.4 


0.8 


9 


5.5 


1.3 


3 


5.2 


0.7 


12 


5.0 


0.0 


2 


5.6 


1.0 


26 


5.4 


0.5 


9 

1 


5.4 


0.5 


10 


5.3 


0.8 


21 



12.1 


0.3 


12.4 


0.5 


12.0 


0.0 


12.0 


0.0 


12.1 


0.3 



12.0 

12.0 
12.3 



0.0 

0.0 
0.5 



Table 10 Frequency distributions of position of first ctenoid scale (relative to anterior lateral line scales) for Anisochromis species. 



/ 



3 



7 



8 



10 



11 



12 



13 



14 



SD 



4. kenyae 
















Kenya 


- 


1 


7 


5 


- 


- 


- 


Mozambique 


2 


1 


3 


2 


- 


- 


- 


Comoros 


- 


- 


9 


7 


4 


- 


2 


Madagascar 


1 


- 


1 


- 


2 


- 


- 


Total 


3 


2 


20 


14 


6 


- 


2 


4. mascarenensis 
















Reunion 


- 


- 


4 


8 


3 


1 


1 


Mauritius 


- 


- 


- 


1 


- 


- 


- 


Total 


- 


- 


4 


9 


3 


1 


1 


4. straussi 


- 




1 


11 


9 


7 


8 



3.3 


0.6 


2.6 


1.2 


4.0 


1.2 


3.5 


1.9 


3.6 


1.2 



4.2 

4.2 
6.7 



I.I 
2.3 



REVISION OF ANISOCHROMINAE 207 

Table 11 Frequency distributions of counts of posterior interorbital pores for Anisochromis species. 

12 3 4 5 x SD 

A. kenyae 

Kenya 

Mozambique - 

Comoros 1 

Madagascar - 

Total 1 

A. mascarenensis 

Reunion - 

Mauritius - 

Total 
A. straussi 4 



3 


5 


2 


- 


2.9 


0.7 


4 


- 


1 


- 


2.4 


0.9 


8 
1 
6 


- 


2 


1 


2.5 


1.2 


5 


5 


1 


2.6 


1.0 


- 


7 
1 
8 


2 


- 


3.2 


0.4 


_ 


2 


_ 


3.2 


0.4 


.5 


- 


- 


- 


1.9 


0.4 



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CONTENTS 



109 Type material of Stegocephalidae Dana, 1855 (Crustacea, Amphipoda) in the collections of 

The Natural History Museum, London, including the description of seven new species 

J. Berge, W. Vaderand A. Galan 
137 The genus Ischioscia Verhoeff, 1928 in Venezuela, with the description of six new species 

(Crustacea, Oniscidea, Philosciidae) 

Andreas Leistikow 
169 A review of the genus Erenna Bedot, 1904 (Siphonophora, Physonectae) 

RR. Pugh 
1 83 A new species of loach, genus Nemacheilus (Osteichthyes, Balitoridae) from Aceh, Sumatra, 

Indonesia 

R.K. Hadiaty and D.J. Siebert 
191 Revision of the western Indian Ocean fish subfamily Anisochrominae (Perciformes, 

Pseudochromidae) 

A.C. Gill and R. Fricke 



ZOOLOGY SERIES 

Vol. 67, No. 2, November 2001